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PROCEEDINGS
OF THE
GENERAL MEETINGS FOR SCIENTIFIC BUSINESS
OF THE
AVOLOGICAL SOCIETY
OF LONDON
1909, pp. 1-544.
(JANUARY—APRIL,)
PRINTED FOR THE SOCIETY,
AND SOLD AT THEIR HOUSE IN HANOVER SQUARE.
LONDCN:
MESSRS. LONGMANS, GREEN, AND CoO.,
PATERNOSTER ROW.
_ 2\2zb5s
by Wsisye de
OF THE
COUNCIL
AND OFFICERS
OF THE
ZOOLOGICAL
SOCIETY OF LONDON.
1909.
COUNCIL.
His Grace THe Duxkt or Beprorp, K.G., President.
GeorGE A. BouLenGcER, Esq.,
F.RB.S., Vice-President.
Prof. J. Rosr Braprorp, M.D.,
D.Sc., F.R.S., Vice-President.
Lt.-Col. Sir R. HAveEtocK-
Cuartes, K.C.V.O., M.D.
ALFRED H. Cocks, Esq., M.A.
Tue Rr. Hon. THE Haru or
Cromer, P.C., G.C.B.
CHARLES DRUMMOND,
Treasurer.
FREDERICK GILLETT, Esq.
F. Du Cane Gopman,
DICA SEERA Ss
dent.
THE Marquis or HAMILTOoN,
M.P.
Esq.,
Ksq.,
Vice-Presi-
SypNey F. Harmer, Hsq.,M.A.,
FE.R.S., Vice-President.
Sir Epmunp G. Lopmr, Bt.
| E. G. B. MrapE-Wa po, Esq.
| Pror.
Epwarp A. MIncuHin,
M.A., Vice-President.
P. CHALMERS MircHety, Esq ,
M-A., DiSc., Tins): Hakuee
Secretary.
W. R. Ocitvre-Grant, Esq.
Apert Pam, Esq.
OLDFIELD THOMAS,
E.R.S.
A. Trevor-Barrys, Esq., M.A.
Epwarp G. WappiLoye, Esq.
A. Smita Woopwarp, Ksq.,
LL.D.,F.R.S., Vice-President.
Ksq.,
PRINCIPAL OFFICERS.
P. Cuatmers Mircnentt, M.A., D.Sc., LL.D., F.BS.,
Secretary.
Frank E. Bepparp, M.A., F.R.S8., Prosector.
R. I. Pococr, F.L.S., Curator of Mammals and Reptiles, and
Resident Superintendent of the Gardens.
D. Seru-Smivu, Curator of Birds and Inspector of Works.
Henry G. Puimmer, M.R.C.S., Pathologist.
F. H. Waternouse, Librarian.
JoHN Barrow, Accountant.
W. H. Coun, Chief Clerk.
LIST OF CONTENTS.
1909, pp. 1-544.
January 12, 1909.
Page
The Secretary. Report on the Additions to the Society’s
Menagerie during the month of December 1908 ......... 1
Dr. H. G. Plimmer, F.L.S., F.Z.8. Exhibition of the
stomach of a Boa suffering from gastritis, and the
stomach and intestines of a Cobra suffering from gastro-
CMG OTD IS ota pcereer corse onrararae = eens uLaisy es tiga wee See oO REE
1. Observations on the Flagellates Parasitic in the Blood of
Freshwater Fishes. By Prof. E. A. Mincuin, M.A.,
VME ZASey U(labesi NE yess ots a siento sei estjages secenee tis
i)
bo
. Zoological Results of the Third Tanganyika Expedition,
conducted by Dr. W. A. Cunnington, F.Z.8., 1904—
1905.— Report on the Copepoda. By Prof. G. O. Sars, —
CAMEZ ES ean (Tz lattes pVell XO Na IMM eee teisecec scene er aise isi 31
. A further Note on the Gonadial Grooves of a Medusa,
Aurelia aurita. By T. Goopry, B.Sc., University
Scholar, Zoological Laboratory, University of Birming-
hams (Plate 2x) es mene aman 2 ied apa, 78
(Ju)
4. The Tuberculin Test in Monkeys: with Notes on the
Temperature of Mammals. By Artaur Erwin Brown,
D.Sc., C.M.Z.8., Secretary of the Zoological Society of
Philagelplia:. s 2... 5. daceae eee seco OR ene eee 81
lv
5. A few Notes on the Whale Balena glacialis and its
Capture in Recent Years in the North Atlantic by Nor-
wegian Whalers. By Prof. R. Couuerr, F.M.ZS.
(Plates XOX XX IT) oe cea cecheeer merrier
February 2,:1909.
Mr. C. Tate Regan, M.A., F.Z.S. Exhibition of two species
of Char (Salvelinus grayt and S. mawillaris) ...............
Mr. R. E. Holding. Exhibition of, and remarks upon, skulls
and photographs of the St. Kilda or Hebridean Four-
horned Sheep, si. .sssssa. ainceeae eee Meer ete see eee eee
Mr. Maleolm Maclaren. Aceount of a fight between a Whale
DMA, SWOLUSH ST: sik ee pees ee ee ee ROE:
Dr. C. W. Andrews, F.R.S., F.Z.S. seeouInN of his visit to
Christmas Island in 1908 =r aCe Tse eA EUR U2.
1. Preliminary Account of the Life-history of the Leaf-
Insect, Phylliwm crurifolium Serville. By H.S. Lerten,
Honorary Research Fellow in the University of Man-
chester, (Plate 2xOxev ail) To eyaceccctuice ve me meeer ne seers
2. The Mammals of Matabeleland. By E. C. Cuuss, F.Z.S.,
Acting Curator of the Rhodesia Museum
3. Report on Deaths which occurred in the Zoological Gardens
during 1908. By H. G. Puimer, F.LS., F.Z.8., Patho-
logist to the Societiyp..-.ccessmte dees set eerie erecta
February 16, 1909.
The Secretary. Report on the Additions to the Society’s
Menagerie during the month of January 1909
Mr. C. Tate Regan, M.A., F.Z.S. Exhibition of sketches of,
and remarks upon colour-changes in some Fishes in
the New York Aquarium
Page
91
98
98
100
101
103
113
—
bo
Or
Vv
Mr. E. G. B. Meade-Waldo, F.Z.S. Remarks upon a letter
from Dr. Einar Loénnberg on the hunting of the Sea-
lephant ony SowthyG cori es.cas sce teaser. cece:
i. The Fauna of the Cocos-Keeling Atoll, collected by
F. Wood Jones. By F. Woop Jones, B.Sc., F.Z.5.,
with the assistance of other Authors ....................6++:
2. Contributions to the Anatomy of certain Ungulata, in-
cluding Vapirus, Hyrax, and Antilocapra. By Frank EK.
Bepparp, M.A., F.R.S., F.Z.8., Prosector to the Society.
3. Le Rhinocéros blane du Soudan (Zhinoceros simus cottont).
Par le Dr. E. L. Trovessart, C.M.Z.S., Professeur au
Muséum d’Histoire Naturelle de Paris. (Plates XXIX.—
DX OSOXI Te) WA Soe RR DME ERER CRP O ppm gnatchBo acc Los tc osocp po cokunacED
March 2, 1909.
Mr. R. H. Burne, F.Z.8. Exhibition of, and remarks upon,
certain elastic mechanisms in Fishes and Reptiles ......
Dr. R. F. Scharff, B.Sc., F.Z.S. Exhibition of some long-
bones and antlers of Reindeer from a cave in County
(Clorplkeistiatd gelichave aera er eee eee eae haa kG oounnate
My. R. I. Pocock, F.L.S., F.Z.8. On the Skulls of Leopards.
pe
. The Development of the Subdivisions of the Pleuro-peri-
toneal Cavity in Birds. By Mare@arer PooLe
bo
. The Growth of the Shell of Patella vulgata L. By E.S.
RUUSSHLL,) MAW? \(Blatie) XOXOXGIEI Rasen. ssetia te seine «2
3. The Life-History of the Agrionid Dragonfly. By Frank
Baurour - Browne, M.A. (Oxon.), F.R.S.E., F.Z.8.
(Plates) XEXOXCINDS de OXOXG NCTA) ieee teased areas eae
i
. Growth-Stages in the British Species of the Coral Genus
Parasmilia. By W. D. Lane, M.A., F.Z.S8., British
Museum (Nat. Hist.)
eee weer eee esse soe eee re eresceeesevesaeesase
Page
132
160
198
201
210
253
March 16, 1909.
The Secretary. Report on the Additions to the Society’s
Menagerie during the month of February 1909 .........
Mr. E. C. Chubb, F.Z.S. Exhibition of the skins and skulls
Of two-foctal LiOns..5..nqlhene eee eee aoe ee een
The Secretary. Exhibition of a photograph of a young
American Waypiy occ cnig.ere acces Mesut eeem emer as eer
The Secretary. Note on a paper by C. Onelli in the
‘Revista del Jardin Zoologico de Buenos Aires’ .........
The Secretary. Exhibition of a photograph of a small herd
of Mountain Zebras (Hquus zebra) ...............c eee eee ee
The Secretary. Exhibition of a photograph of a female
Giraffe captured in West Soudan ......1.........0¢-:--e0e-:
Dr. F. A. Bather, M.A., F.Z.S. Exhibition of a fossil
Hehinoidu Seucellanainateia epee. coe eee ee
1. The Kctoparasites of the Red Grouse (Lagopus scoticus).
By A. E. Surptry, M.A., Hon. D.Sc., F.R.S., F.Z.S.,
Fellow and Tutor of Christ’s College, Cambridge, and
Reader in Zoology in the University. (Plates XXX V.—
OXI VT.) oc ee ee etna a tece eee on aeons ame ces meenecier
2. The Thread-Worms (WVematoda) of the Red Grouse
(Lagopus scoticus). By A. HK. Surpuey, M.A., Hon.D.Sc.,
F.R.S., F.Z.8., Fellow and Tutor of Christ’s College,
Cambridge, and Reader in Zoology in the University.
(Plates XOn VEE SIV) eee acest chro ones eee eee vane
3. The Tape-Worms (Cestoda) of the Red Grouse (Lagopus
scoticus). By A. E. Surpiey, M.A., Hon. D.Sc., F.R.S.,
F.Z.8., Fellow and Tutor of Christ’s College, Cambridge,
and Reader in Zoology in the University. With a Note
by Wm. Byerave, M.A. (Plates LVI.-LX.) ............ 5
Page
308
308
308
308
308
509
309
309
33D
Vil
Page
4, Internal Parasites of Birds allied to the Grouse. By A. E. :
SareLey, M.A., Hon. D.Sc., F.R.S., F.Z.S., Fellow and
Tutor of Christ’s College, Cambridge, and Reader in
Aoolosyrinathe Winthyersiuygmeer eres sreesass ase neee - 363
5. On a Fossil Bird from the Lower Pliocene. By W. P.
BVI CR AMIN GHZ (150, JAC aS oo rere nmr ee rere le a cee eet OLE cas 368
6. On a Collection of Mammals from Western Java presented
to the National Museum by Mr. W. E. Balston. By
OLDFIELD Tuomas, F.R.S., F.Z.S., and R. C. Wroucuton,
IE) ZI Bu Poss ded's'o cic oS unk A Rey ata ae ORE Rina ete re ron 371
April 6, 1909.
Mr. George Jennison. Exhibition of, and remarks upon,
some fertilized eggs from a pair of Seba Pythons......... 392
Dr. R.T. Leiper, F.Z.8. Exhibition of a distorted Elephant’s
tusk and a malformed canine tooth of a Hippopotamus . 393
Mr. E. T. Newton, F.R.S., F.Z.S. Exhibition of a meta-
tarsal bone of an Ox showing the marks of gnawing by
POGOIUSH is cheats ee, cae aia esertais Seas eate ee Sane ee tee ee 393
1. Description of a new Form of Ratel (JWellivora) from
Sierra Leone, with Notes upon the described African
Forms of this Genus. By R. I. Pocock, F.LS., F.Z.S.,
Superintendent of the Gardens. (Plate LXI.) ......... 394
2. Notes on an Ichthyosporidian causing a Fatal Disease in
Sea-Trout. By Muriet Roperrson, Carnegie Research
Helllow..) (Plates TiXel—1iXelaye ere pears eee ee) 399
3. A Collection of Fishes made by Dr. C. W. Andrews, F.R.S.,
at Christmas Island. By C. Tarr Ruaan, M.A., F.Z.S.
(Gedy enim DD: Oh Priced De. @ CS) Be So nonortocednccenaodousoatcnneateee ee 403,
4, On some New and Little-known Hesperiide from Tro-
pical West Africa. By Hamitrton H. Drucs, F.LS.,
BAL Staal (Eta tee aXe Valsle aye eaene ae ere nee nee hoses, 406
vill
April 27, 1909.
The Secretary. Report on the Additions to the Society’s
Menagerie during the month of March 1909
Seer eee eee ece
Mr. H. F. McShane. Exhibition of a series of lantern-slides
of animals living in the Society’s Gardens
eee e sc ero eee se seee
Prof. E. A. Minchin, M.A., V.P.Z.S. Exhibition of the
eggs, and liying specimens of the larve and adults of the
Tick, Ornithodoros monbata Murray
CO iO a i ie icin arary
Mr. R. H. Burne, F.Z.8. Exhibition of a series of specimens
showing the Anatomical mechanisms of the eyes of some
Vertebrates
See meee cere ere cence essere rere ssceceeseeneeeceseseerecce
Mr. R. I. Pocock, F.L.8., F.Z.S. Exhibition of, and remarks
upon, the photographs of two Quaggas, or Burchell’s
Zebras
Coe ee nese eeereaecresreseesscree seer ease eeee eee np tee esseeeeesoen
1. A Review of the Species of the Lepidopterous Genus
Lycenopsis Feld. (Cyaniris auct. nee Dalm.) on exam-
ination of the Male Ancillary Appendages. By T.
ANMCIDEANONY (CIEUNEWUNIE, IWEID) IN WAISS 6 inks stn shen soocodosectac
bS
. On some Points in the Structure of Galidia elegans, and
on the Posteaval Vein in Carnivores. By Frank E.
Bepparp, M.A., F.R.S., F.Z.S., Prosector to the Society.
3. On the Postcaval Vein and its Branches in certain
Mammals, By Franx E. Bepparp, M.A., F.R.S., F.Z.S8.,
Prosector to the OGLE bYy. GMs Gere cme e rere etic he ae
4. On the Comparative Osteology of the Passerine Bird
Arachnothera magna. By R. W. Suuretpr, M.D.,
CAVMEZ ES t( Plate {aay aA Te acces eee eee er eee,
Page
414
414
4}4
419
477
496
ALPHABETICAL LIST
CONS a Ol ORs:
With References to the several Articles contributed by each.
(1909, pp. 1-554.)
Page
ANDREWS, CHARLES WiturAM, B.A., D.Sc., F.R.S., F.Z.S.
Account of his visit to Christmas Island in 1908 ...... 101
Baurour-Browne, Frank, M.A. (Oxon.), F.R.S.E., F.Z.8.
The Life-History of the Agrionid Dragonfly. (Plates
DO. GT ET: OO. G IG erence erro iecteede aco nacaosde 253
Baruer, Dr. Francis A., M.A., F.R.S., F.Z8.
Exhibition of a fossil Echinoid, Sewtellina patella ...... 309
BeppDarD, Frank EH., M.A., F.R.S., F.Z.8., Prosector to the
Society.
Contributions to the Anatomy of certain Ungulata,
including Tapirus, Hyrax, and Antilocapra ...........065- 160
On some Points in the Structure of Galidia elegans,
and on the Postcaval Vein in Carnivores ................6+ AT7
On the Postcaval Vein and its Branches in certain
BU ewan ated Ship ee Cs te al Marl fe cin td ltl ha A eaten ed can an 496
ae
x
Page
Brown, Artuur Erwin, D.Sc., C.M.Z.S., Secretary of the
Zoological Society of Philadelphia.
The Tuberculin Test in Monkeys: with Notes on the
Temperature of Mammals... .as-)seusocceeeeeeee eee ees 81
Browne, Frank BAatrour-. See BALFouR-BRrowne, FRANK.
Burne, RicHarpD Hicerns, F.Z.8.
Exhibition of, and Remarks upon, certain - elastic
mechanisms in Fishes and Reptiles ...............-...-....--- 201
Exhibition of a series of specimens showing the
Anatomical mechanisms of the eyes of some Vertebrates. 414
Byerave, Wm., M.A. See Surptey, ArtHUR EVERETT.
The Tape- Worms (Cestoda) of the Red Grouse (Lagopus
scoticus).
CuHapMAN, T. AugEeRNON, M.D., F.Z.S.
A Review of the Species of the Lepidopterous Genus
Lycenopsis Feld. (Cyaniris auct. nec Dalm.) on exam-
ination of the Male Ancillary Appendages .................. 419
Cuuss, Ernest Caries, F.Z.8., Acting Curator of the
Rhodesia Museum.
hiemViammalsiorVlatalbele! ancl sss eee nee rer Hales
Exhibition of the skins and skulls of two feetal lions... 308.
Couuert, Prof. Ropert, F.M.Z.S.
A few Notes on the Whale Balena glacialis and its
Capture in Recent Years in the North Atlantic by
Norwegian Whalers. (Plates XXV.-XXVIL.)............ 91
Xl
Page
Druce, Hamiuron H., F.L.S., F.Z.S.
On some New and Little-known Hesperiid~ from
Rgppreall West Atrica), 9) (a laibe li XGVally see sas: 5a s2.455 2. 406,
Goobzy, T., B.Sc., University Scholar, Zoological Labor-
atory, University of Birmingham.
A further Note on the Gonadial Grooves of a Medusa,
FNOTRGINO) CDH Oi A VARI) DOSOINYS) ooucode Gososoboacudcoacaason 78
Hotoine, R. E.
Exhibition of, and remarks upon, skulls and photo-
graphs of the St. Kilda or Hebridean Four-horned
INCE Os Se evacie ett cae o sisicii acs aicorarasteletiats => ene ROSE A TERRE RIS sreeee 98
JENNISON, GEORGE.
Exhibition of, and remarks upon, some fertilized eggs
Lrommaypale ob Seba k ybhOusy.c-a-.cueh we cee een ee eet ee 392.
JONES, FREDERICK Woop, B.Sc., F.Z.8., with the assistance
of other Authors.
The Fauna of the Cocos-Keeling Atoll, collected by
1 AW oo dl Ke) 02 ae REE Ree Gi » scrHSaatnnduddetacchicHot 132
Lane, Witiiam D., M.A., F.Z.S., British Museum (Nat.
Hist.).
Growth-Stages in the British Species of the Coral
Genus Pargsnvilidr .i2c4bich ese eeeee ee ee 285
Leicu, H. 8., Honorary Research Fellow in the University
of Manchester.
Preliminary Account of the Life-nistory of the Leaf-
Insect, Phyllium crurifoliwm Serville. (Plate XXVIII.) 103
Page
Leirrer, Dr. Roser T., F.Z.8. 4
Exhibition of a distorted Elephant’s tusk and a mal-
formed canine tooth of a Hippopotamus .................+0.+ 093
McSuane, H. F.
Exhibition of a series of lantern-slides of animals living
Tal (Hate) OV Uys) CRMC | Sac geronasseosononacaoouosnusensoodsacs 414
MacLAREN, MAtcotm
Account of a fight between a Whale and a Sword-fish. 100
Meave-Watpo, E. G. B., F.Z.S,
Remarks upon a letter from Dr. Einar Lénnberg on
the hunting of the Sea-Elephant on South Georgia ...... 132
Minouin, Prof. Epwarp A., M.A., V.P.Z.S.
Observations on the Flagellates Parasitic in the Blood
of Freshwater Fishes. (Plates I1—-V.) ..............0000..5 2
Exhibition of the eggs, and living specimens of the
larvee and adults of the Tick, Ornithodoros monbata
Miuaeraiy "1, dic eee eto ne at operas 2ke en Ce AEA At a Al4
Mitcnett, P. CHatmers, M.A., D.Sc., LL.D., F.RB.S.,
F.Z.S., Secretary to the Society.
Report on the Additions to the Society’s Menagerie
during the month of December 1908 ..............0..200005 1
Report on the Additions to the Society’s Menagerie
duane ice) mont hyo ama ty 200 09 rene nee eee eee ete ae 130
Report on the Additions to the Society’s Menagerie
during the month of February 1909
Exhibition of a photograph of a young American Tapir. 308
Note on a paper by C. Onelli in the ‘ Revista del Jardin
ZoolosrcondesBuenos Aires a sa ee eee eee ee 308
Xl
MircHeny, P. Cuaumers. (Continued.)
Exhibition of a photograph of a small herd of Mountain
Lie wea (Hi 1s 2CO TG) Nene eet ees teal ack ssaie aioe sce v0 a
Exhibition of a photograph of a female Giraffe captured
in West Soudan
woe eet ewe ere eee meet wees e eee eer eeeroesesreneeeesnene
Report on the Additions to the Society's Menagerie
during the month of March 1909
eee eet eee e trees ee ee eee eeeres
Newvon, Epwin '., F.R.S., F.Z.S.
Exhibition of a metatarsal bone of an Ox showing the
MarkslOh enawane by, rOd entisi yas peeeete eee ee ner teen
Purmmer, Henry G., M.R.CS., F.L.S:, F.Z.8., Pathologist
to the Society.
Exhibition of the stomach of a Boa suffering from
gastritis, and the stomach and intestines of a Cobra
suffering from gastro-enteritis
Ci SC eee ae a en i Ca
Report on Deaths which occurred in the Zoological
Gardens during 1908
Pocock, Rreainaup I., F.L.S., F.Z.8., Superintendent of the
Gardens.
Onithe: Skulls'ot MWeopards sess ere coerce eee
Description of a new Form of Ratel (Jellivora) from
Sierra Leone, with Notes upon the described African
Hormeror this Genucy | (later axes ieee ee tenes eee aes
VY Exhibition of, and remarks upon, the photographs of
two Quageas or Burehell’s| Zebrasy (7... steers - secre... << ers
Poouz, Miss MARGARET.
The Development of the Subdivisions of the Pleuro-
peritoneal Cavity in Birds ....-..0secseceeeee sec ctececeeer eens
308
393
204
394
X1V
Page
Pycrarr, WiLirAM P., F.Z.8., A.L.S.
On a Fossil Bird from the Lower Pliocene ............... 368
Reaan, C. Tarn, M.A., F.Z8.
Exhibition of two species of Char (Salvelinus grayi
amd ySe acc ularts) 240% vseseacecewnctoce nee meee Ce ee seer ee ee 98
Exhibition of sketches of, and remarks upon colour-
changes in some Fishes in the New York Aquarium ... 130
A Collection of .Fishes made by Dr. C. W. Andrews,
F.R.S., at Christmas Island. (Plates LXV. & LXVI.). 403
Ropertson, Miss Murreu, Carnegie Research Fellow.
Notes on an Ichthyosporidian causing a_ Fatal
Disease in Sea-Trout. (Plates LXIT.—LXIV.) ............ 399
RussEwx, BE. 8., M.A.
The Growth of the Shell of Patella vulgata L. (Plate
DON Ta) diac huis. saceetn ectiee Seite os eae oils asin pis ceetnas ac amram ate 235
Sars, Prof. Grorce O., C.M.Z.S.
Zoological Results of the Third Tanganyika Expedition,
conducted by Dr. W. A. Cunnington, F.Z.8., 1904-
1905.—Report on the Copepoda. (Plates VI-XXIIT.). 31
ScHarFF, Dr. Roperr F., B.Sc., F.Z.8.
Exhibition of some long-bones and antlers of Reindeer
iromeaicave 1 CountvaCorkarmelinel amd str enpemrece acces eee 204
Suipiey, ArrHuR Everert, M.A., Hon.D.Sc., F.RB.S., F.Z.8.,
Fellow and Tutor of Christ’s College, Cambridge,
and Reader in Zoology in the University.
The Ectoparasites of the Red Grouse (Lagopus scoticus).
(Plates XV OV.) cs. ite eae eae eon areas 309
XV
SarpLey, AnrHur Evererr. (Continued.)
The Thread-Worms (Nematoda) of the Red Grouse
(Lagopus scoticus), (Plates XLVIII.-LV.) ...............
The Tape-Worms (Cestoda) of the Red Grouse (Lago-
pus scoticus). With a Note by Wm. Byerave, M.A.
(GEES ORY IR OD.) Pan tee i nO age lst at enon tna
Internal Parasites of Birds allied to the Grouse ......
SHuFELpr, Roperr W., M.D., C.M.Z.8.
On the Comparative Osteology of the Passerine Bird
Arachnothera magna. (Plate LXVIIT.) ............0.....
Tomas, OLDFIELD, F.R.S., F.Z.S8., and Wrougnton, RoBpert
OR WAI
” On a Collection of Mammals from Western Java pre-
sented to the National Museum by Mr. W. E. Balston...
‘Trovgssart, Dr. Epovarp Louis, C.M.Z.S., Professeur au
Muséum d’Histoire Naturelle de Paris.
Le Rhinocéros blane du Soudan (Zhinoceros simus
COLL Ona aes (llabes) NEXaeXe— XONOXGID) ee eee eee
Watpo, E.G. B Meaps-. See MrapE-WALDpo, E. G. B.
Wrovcuton, Ropert C., F.Z.8., and THomas, OLpFrexp,
F.R.S., F.Z.8.
On a Collection of Mammals from Western Java pre-
sented to the National Museum by Mr. W. E. Balston ...
Page
335
we)
~J
—
198
df]
4
Meg
Ri ba #
M} ee Het til 1 Cy
ear
why:
i. nl
XXiI.
XXII.
XXIII.
LIST OF PLATES.
1909, pp. 1-544.
Page
Trypanosomes of Hel and Perch ....:.--.....-.... >
Trypanosomes of Pike, Tench, and Bream. Trypano-
plasms of Pikes: eacee ak ie elec) eet
Trypanoplasms of Tench, Bream, and Rudd ........ We
somes of Tench
Trypanosomes of Hel and Perch
Diaptomus galebordes G. ©. Sarvs. .-...2..+--20----
Draptoniuis macs: Gi OS aC Sei ein eit dere ere
Diaptomus stuhlmanni Mrazek and D. simplex G.O.Sars
Diaptomus cunningtont G. O. Sars ......... 000000
Schizopena mopnata G. OF Sars sae ssen ss ee eee |
Schizoperu validior G. O. Sars, S. consimilis G. O. Sars, |
anges. wgulata, Gy OS ats meer lateles eri neer
Schizopera minuticornis G. O. Sars, 8. spinulosa G. O.
Sars, and S, fmobreata’ Ge @Sars2 so. e e adene -
Schizopera scalaris G. O. Sars and Ilyophilus perplexus
Go Ol Sarg: ue.) says si aspeteeean renee sleaaistarsn onsets are
Cyclops leuckarti Claus, C. emint Mrazek, and C. ne-
GUAGE AC OMORSENIS Bo amoonsochcoouodcogueg Sado.
Cyclops tenellus G. O. Sars and C. albsdus (Jurine) .
Cyclops attenuatus (. O. Sars, C. varicans G. O. Sars,
pl (Oho Cm Oh cocsdueonsbonectoocuas
Cyclops cunnington: G. O. Sars and C. pachycomus G.
CRISTO SO Etocoodd shonanooUda Leo
Cyclops semiserratus G. O. Sars ...............40.
Cyclops levimargo G. O. Sars, C. angustus G. O. Sars,
Praal (On puanmeucons (Cr (SEV “Spcccnecos00500004
Cyclops agiloides G. O. Sars, C. ewacanthus G. O, Sars,
and Cy crliaiuseGn OM Sans ae eera ieee sien ye
Cyclops oligarthrus G. O. Sars, C. compactus G. O. Sars,
ands @s dubwus, GO Saisie tee law es snl 5 «
Ergasiloides megacher G. O. Sars ...........0...
Ergasiloides macrodactylus G.O. Sars and £. brevi-
GHG, OSE sooongnocoedo sondoneoecnpbos \
Trypanoplasms of Pike, Tench, and Bream. Trypano- f
31
Proc. Zoou. Soc.—1909. b
XV1ll
Plate Page
XXIV. Gonadial Grooves of Awrelia aurita ....... cee ee 78
XXV,
XXVI. | Balena glanaus 2209. ah eee eee 91
XXVIL. ce) .
ON VME Pay llm)eruriyjolcum pe a eee ae eee 108
XXIX. Rhinoceros sitmus cottont S, DP ...........+-+:-::
coe Rianoeeros simus cotton: 8... .......--+2-+ +++" | 198
XXXII. Shells of Patella vulgata, XZ2 ....-- +1. seve erences 285
era, Development of Agrionid Dragonflies .........--: 253
XXXV. )
- XXXVI ake
MX XVI. -Ganiodes tetraonts, ). oa ja0) Sy leery geet os Wars )
XXXVIL | 7 |
XXXIX. } |
XL. Gontodes tetraons and Menopon pallescens ........+- |
OE (Caves eine l INO P TONS sa ond adacodogs soe 4a4o5 Tee a
GUI Worms eamenateus 20 Wl tay tak oie PE UREEE .. 209
ene Onnithoniyjia (agopodts (na... 6 pear era hae
aoe (Sedéophanarsterconat@ = is ne erry eee * |
XLVII. Ceratophyllus galinule ... 2.0.11 2111s ees J
XLVUI> |
ALIEN \ Trichostrongylus MAPUPUGIUT Vance cion ditn ooo dagbad>
|
Tid habia |
LI. Trichostrongylus pergracitis and T. richosoma longicolle . L335
LUT. | Trichosoma Longicolles Ph 207.2). ieee Wee acest eee :
LIV. { ;
LV. Contents of alimentary canal of Grouse .........:.. J
LVI, Davamen urogalli ee ene Peer i rs
LVIL. Davainea urogalli and Hymenoleps MICPOPS .... apceall
LVIII.. Hymenolepis microps ...:.-....++-. Tacs! im boonte oye 351
IDX, Davanea an ogollt Gere ere eee ee eee eee |
LX. Davainea urogalli and Hymenolepis microps ........ Pa
TUXGy Mellsvorasignate soe. nee ia) eee eee hs re oot
LXIL } -:
LXII. fJouyonesiion of Sea-Trout .......... mencgege 399
LXIV.
INOW 1) Parupeneus andre weiter. © 1h (laine eter eee >
LXVI. 1. Blennius atrocinctus. 2. B. nativitatis. 3. Salarias |,
caudofasciatus. 4. S. natalis, 5, S. melanosoma. (ae
Te (OM AACHEAS TMU. Bacdgogbocoxdsos555G800 gr)
LXVIL. New and Little known W. African Hespervide...... 406
LXVIIL. Skeleton of Arachnothera magna ...... 0... cree ~ 527
Ik
LIST OF TEXT-FIGURES.
1909, pp. 1-544.
Page
Cercopithecus pygerythrus 3 adult. Temperature chart showing
normat daily curve ...-...... wh cba staat Rare Peps mer ee on re Perrmitc |
Macacus rhesus § juv. T sxpemsinne chart thane ater
culous. reaction destroying daily curve, with no general
rise. Ai HbR Rn ere IOP GO ooo. GOS 6 om OOo.G0 pk Hee Ce oo sao &4
Cer copious ee 2 adult. Temperature aba showing
marked post-injection fall in advanced tuberculosis ........ 85
Lemur varius 2 adult. Non-tuberculous temperature charts 87
Balena glacialis, male. ea eas Oretaeere SER ee a) are Up Wee OS
Heads ete. of Four- horned Sheep) «kee aoe ye eae oS)
Eee of Gygis candida, laid on the heel of a grongong tree
(Cordia subcordata)... BECO OMT ODNe GOB ie eo bo oT MIRC Gates
Gygis candida sitting on As OOO honevn dap aenar alae Seameel neuer ea h , 39
WONG? CHIGOS COMI Soocooban tn beenecacucgucucor F 142
Lateral view of the Sylvian region of aleve tine ae iipnae
COPICNSIS Hd -neh sete MES CNS Bre eee A in 0 or ein Senta here re etera KOT)
Brain of adult Pao ul: (Antilocapra americana), dorsal
WAOW Sirsa 4 tag ooo Nee eee TORI COO MR pee Ienee & 173
Antilccapra americana. Musculature of outside of forearm.... 178
. The colic helicine in various Artiodactyles ...,...... omnes 183
Ceecum, colic helicine, and adjacent parts of the gut of 1 Paqulits
stanleyanus .,.,.,+: BE ESE a ORISn Bil Ouchy See ee ee 184
xx
. Ceecum, colic helicine, and adjacent parts of the gut of Moschus
TEO/(AUS. soonnscn cou0D ML NRRCH EC OG-0 0.6 Jo moB Ob
. Brain of Madoqua phillipsi, dorsal view .........+.....0000-
. Brain of Madoqua phillips, lateral view ............++..
. Brain of Babyrussa alfurus, dorsal view ...-.....+....++..+-
. Brain of Babyrussa alfurus, lateral view ..........+.....----
. A transverse section through the vertebral column and aorta
Page
of a Shad, in the region of the trunk ...... we lala dee Satay eo%e 202
. Diagrams of one and the same portion of the aorta, showing
the relative positions of the ventral ligament and aortic cavity
during the lateral flexions of the body in swimming........ 2
Skull of Leopard (Felis pardus oe from Cette
OnviaEMe seb onaes cob aob ua GQonododdonoUAmUsoo GODS
Skull of Leopard (Fels pardus pares from NIiow, CP,
Tn taints. tatekare et Me PR rte c's as Oe ae ate ah
. Transverse section of a chick of 5 days’ incubation, Dennen he
anterior region of the liver; anterior face ................
. Slightly diagrammatic reconstruction from horizontal longi-
tudinal sections of a chick of 6 days’ incubation ; seen from
the "ventral Side sgs satan wise esc os oe eens Sen Nae ole rh a ene
Transverse section of a chick of 7 days’ incubation, through the
posterior part of the pericardium and anterior part of the
liver; anterior face ..... Sea eer: EN SOR MTL A ERS ea Sir
. Longitudinal vertical section of a chick of 7 days’ incubation,
taken a little to the left of the middle line; seen from the
OTM OM awh code dodasds wo dIDodocUDe DUC Dogar Secon
28. Reconstruction, similar to text-fig. 25, of a chick of 8 days’
MNGMISEMNCIIG oa ccgohangaseguac seal des od Have ahaa suataldeene es
29, Reconstruction, similar to text-figs. 25 and 28, of a onel of 10
GERGE? HACKRTNCN ogocn09ggs0ce0copasguc 99 Geos DOL acces
51.
. Transverse section of a chick of 13 days’ incubation, Hhroueh the
region of the pericardium and anterior intermediate air-sacs ;
seen from behind....... SR Eh Nike aces Pee Rhea we Pee
Transverse section of a chick of 13 days’ incubation, through the
connection between the posterior part of the post-pulmonary
septum and the mesentery of the liver and gizzard; seen from
TEI hous ae cna coas-ae-s0 6 adeeiatety HOS PA ;
Adult Rook bisected horizontally through the region of The
gizzard and liver-lobes, and slightly dissected attenionly to
expose the lungs; seen from the ventral side........ Baie bio
. Diagrammatic plan showing the subdivisions of the coelom in a
bird in longitudinal section ...... Bri oes apices hg ene Seek
. Chick of 17 days’ incubation bisected longitudinally a little to
the left of the middle line; seen from the right side ......
. Young Crocodile bisected in the same way as Chick in text-fig. 34 2
218
214
219
291
XX1
Page
36. Young Crocodile opened from the ventral side ............ 231
37. Diagrammatic plan showing the subdivisions of the ecelom in
a Crocodile im longitudinal) section 9). .2.). Wee 65.0.5. 232
38. Diagram illustrating the progress of costal ornamentation in
the English species of Parasmilia.......06.e. sce e encase 289
39. Costal stages in P. serpentina Kdwards & Haime..........., 2938
AOMCostallstages| in’ Pi centralis Memtelly ete tae eee se. 295
41. Costal stages in P. fittont Edwards & Haime................ 296
49. Costal stages in P. granulata Duncan ............ cesses ees 298
43. Costal stages in P. gravest Kdwards & Haime .............. 300
44, Costal stages in P. mantelli Kdwards & Haime ............ 302
45. Costal stages in P. cylindrica Edwards & Haime............ 303
AGw Costalystaces un i. mores Wun aiieratyetetey- = eyayalel-vaiete steele tt 305
47. The Slab and the counterpart thereof containing remains of a
Morssill Lerqamns (lau anIg WORTOUEEIHD)) 6.0 poe ob on Seog noengeoDuECaS 369
48. Burchell’s Quagga (Equus quagga burchelli) ...... ccc eevee 415
49 Wahlberg’s Quagga (Hquus quagga wahlbergt) .....1-....+e. 416
50. Diagram of one of the northern races of Equus quagga ...... 418
51-56. Ancillary Appendages of species of Notarthrinus .... 424-427
57-117. Ancillary Appendages of species of Lycenopsis .... 428-472
118. Ancillary Appendage of Bothrinia nebulosa ........0..eee-- 472
119. Ancillary Appendage of Artopéetes pryert .....-......000- 474
120, Ancillary Appendage of Megisba malaya ........+-. 2000 0ee AT4
121. Ancillary Appendage of Neopithecops zalmora .............. 475
122. Ancillary Appendage of Castalius ethion......-...00.ssee ees 475
123) Palate of Galidia clegans 1a. we «ce ele elm ee = oe oleae = 478
124, Tongue of Galidia elegans, dorsal view ........++++.-+++0-- 479
125. Caudate lobe of liver, duodenum, and adjacent structures in
Gahdia elegans........+ AR ao a MIC MILO olo Clos 48]
126. Cxecum and adjacent structures of Galdia elegans .......... 482
127. Liver of Galidia elegans, abdominal surface ................ 483
128. Brain of Galidia elegans, dorsal and lateral views .......... 484
129. Postcaval vein and branches in Galidia elegans and inept
WOE Ron peoe mene chon oubocOocccodpccocsaroseeoooar 486
130. Posteaval vein and branches in Suricata tetradactyla and
Cercoleptes caudivoluulus. .....ssccacsneecncseescensceees 488
131. Postecaval vein and branches in Melhvora signata, Ictonyx
capensis, and Mephitis mephitica ......-.00. seve eee eee 491
. Posteaval vein and its branches in T7ichosw'us vulpecula and
GUNG SOTRUES | OUMOUIODSOE) 50 Oma 00000 oc0s50050 bo ceobnnGOe 499
3. Postcaval vein and its branches in Dasyurus maugei and
Onychogale frend... ce cree cece ener ce cee e ete e nena 502
. Posteaval vein and its branches in Macropus parryi and
WER HIS CUS. Gocosocoodueccdcouo tbmabseoouCONDObEEE 504
. Postcaval vein and its branches in Zatusia peba and Dasypus
PEMETOSUS! cesiadnlorelcunce esi pisediorare: Geers FA NCROR LAURER GRA CRON CIES soan BOs)
XX11
Page
136. Posteayal yein and its branches in Centetes ecaudatus and
Lysiurus unicinctus ...... ih cick akihe mele ee Pep uiele ¢ 508
157. Posteaval vein and its branches in Merocebus smithid and
Lemur catta ..... Sars ack bac ‘ see cepee clk bh aMOle
138. Postcaval vein and its branches in sean Us pieces Th anomie
chrysogaster, and Dipus hirtipes ..... a Ao Shean ine! Sole
159, Postcaval vein and its branches in Zapus aon Gerbillus
egyptius, and Geor hychus capensis FE Sees AIRE nee ee ns We 519
LIST OF NEW GENERIC TERMS
PROPOSED IN THE PRESENT VOLUME (pp. 1-544).
Page Page
Ariopoetes (Lepidopt.) ......-.. 473 Ergasiloides (Copepoda) ...... 63
475
Bothrinia (Lepidopt.) .........
“ee
.
PROCEEDINGS
OF THE
GENERAL MEETINGS FOR SCIENTIFIC BUSINESS
OF THE
ZOOLOGICAL SOCIETY
OF LONDON.
1909.
Pages 1-200.
Part I. CONTAINING PAPERS READ IN
JANUARY ann FEBRUARY.
JUNE 1909.
PRINTED FOR THE SOCIETY,
SOLD AT THEIR HOUSE IN HANOVER SQUARE.
LONDON :
MESSRS. LONGMANS, GREEN, AND CO,
PATERNOSTHR ROW,
[Price Twelve Shillings. |
a ES
LIST OF CONTENTS.
1909, pp. 1-200.
January 12, 1909.
: Page
The Secretary. Report on Additions to the Society’s Menagerie during the month of
Mecemiber [GOS Giscl sals's atel win teinern wcvelevevs lois keleyete tare lo steleeya ete she pe lade tenetel aie temttede ee sieer rae!
Dr. H. G. Plimmer, F.L.S., F.Z.S. Exhibition of the stomach of a Boa suffering from
gastritis, and the stomach and intestines of a Cobra suffering from gastro-enteritis.... 1
1. Observations on the Flagellates parasitic in the Blood of Freshwater Fishes. By Prof.
HA] Mincrin, MoAs VP ZiS.) (Plates Ve)i rete enelerereiolotel sey eiatote els olen slelerenetaateteteete 2
2, Zoological Results of the Third Tanganyika Expedition, conducted by Dr. W. A.
Cunnington, F.Z.S., 1904-1905.—Report on the Copepoda. By Prof. G. O. Sars,
C.M.Z.S. (Plates VI-XXIII.)..........000. alot gis oisieiove of otmreneis tele sieves) eee 31
8. A further Note on the Gonadial Grooves of a Medusa, Aurelia aurita. By T. Goonny,
B.Sc., University Scholar, Zoological Laboratory, University of Birmingham. (Plate
2 OSIM EE SO SRI GIODIOORADMOGDS OA OE BES uinac ss 3 als ote) Suge eetets wir tae etn ee 78
4. The Tuberculin Test in Monkeys: with Notes on the Temperature of Mammals. By.
Artuur Erwin Brown, D.Sc., C.M.Z.8., Secretary of the Zoological Society of Phila-
Gl NNE a5 cacsdauoseddoguaenScHG00nGd5 ag05 00 Hooded ddGe peso Ha GueSCOsD CC seieeeee 81
5. A few Notes on the Whale Balena glacialis and its Capture in Recent Years in the
North Atlantic by Norwegian Whalers. ByProf. R.Cou.ert, F.M.Z.S. (Plates XXV.—
RK VIE) pee Ridius tune ee ‘td Gees tae i ea of Rs ee eae ae HL
Contents continued on page 3 of Wrapper.
PROCEEDINGS
OF THE
GENERAL MEETINGS FOR SCIENTIFIC BUSINESS
OF THE
ZOOLOGICAL SOCIETY OF LONDON.
(January to April, 1909.)
January 12, 1909.
Prof. J. Rose Braprorp, M.D., D.Sc., F.R.8., Vice-President,
in the Chair.
The Secretary read the following report on the additions made to
the Society’s Menagerie during the month of December 1908 :—
The number of registered additions to the Society’s Menagerie
during the month of December last was 57. Of these 36 were
acquired by presentation, 8 by purchase, 7 were received on
deposit, 2 by exchange, and 4 were born in the Gardens. The
number of departures during the same period, by death and
removals, was 212.
Amongst the additions special attention may be directed to :—
A Brown Bear (Ursus arctos), from Russia, presented by
G. Cecil Whitaker, Esq., on December 8th.
Two Mole Rats (Georychus hottentottus), new to the collection,
from Mashonaland, presented by J. ffolliott Darling, Esq., F.Z.S.,
on December 9th.
Two Labrador Jays (Perisoreus atricapillus), from Labrador,
presented by the Rev. W. W. Perrett on December 14th.
One Yellow-billed Sheathbill (Chionis alba), from the Falkland
Islands, presented by F. W. Tode, Esq., on December 14th.
ee
Dr. H. G. Plimmer, F.L.S., F.Z.8., Pathologist to the Society,
exhibited the stomach of a Boa suffering from gastritis, and the
Proc. Zoou. Soc.—1909, No. I. 1
{YS
2 PROF, E, A. MINCHIN ON PROTOZOAN [ Jan. 12,
stomach and intestines of a Cobra suffering from gastro-enteritis,.
and made the following remarks :—
“These specimens are shown in order to call attention to the
condition of inflammation of the stomach and intestines in snakes.
from the Society’s Collection. Out of 160 cases examined during
the last year by far the larger number showed this condition in
varying degr ees; and that is after excluding those cases due to
the direct Suoidniionl of worms which had cemeed ulceration or
perforation.
“Of course there are many causes of inflammatory conditions
of the stomach and intestines, but it would appear, from the large
percentage of cases showing this condition, that there must be
some common cause, and as the present method of feeding the
snakes is an unphysiological one, it might be worth while to
consider whether it may ‘not be the cause of the large mortality
from these inflammatory conditions of the alimentary tract.”
The following papers were read :—
1. Observations on the Flagellates Parasitic in the Blood of
Freshwater Fishes. By Prof. E. A. Mincuty, M.A.,
Wale AS)
[Received December 23, 1908. ]
(Plates I.-V.*)
Introductory.
The trypanosomes and trypanoplasms of freshwater fishes have.
been studied by a number of naturalists abroad, and have been
the subject of several important memoirs, more especially by
Laveran & Mesnil, Léger, and Brumpt in France, and by Keys-
selitz in Germany. In this country, however, little attention has
been paid to them. I was therefore glad to avail myself of the
exceptional opportunities offered by the Sutton Broad Laboratory,
to study the parasites of the fishes in the Norfolk Broad. TI desire
to take this opportunity of expressing my warmest thanks to my
friends Messrs. Kustace and Robert Gurney for their kindness in
putting at my disposal the resources of their picturesque and well-
equipped laboratory, and for much help during my stay there. I
spent portions of my summer vacations at the laboratory, about
three weeks in August and September 1907, and five weeks in the
same months in 1908, and during these periods I occupied myself
almost entirely with these parasites. Although I have only
touched the fringe of the question so far, and there is still every-
thing to be discovered and worked out concerning the all-important
* Wor explanation of the Plates see p. 29.
‘14
M.R.adnat.del. Lith. Anst.v.b A. Punke, Leipzig.
TRYPANOSOME'S OF EEL AND PERCH.
fith. Anst.v BA Funke, Leipziq.
AND BREAM.
M.R adnat del,
TRYPAN OS OME SO Pb yal BANG ii
TRYPANOPLASMS OF PIKE.
1D Wansie JCONS)) SACI NE
MR. adnat del. Lita. Anst.vE.Afunke, Leipzig.
TRYPANOPLASMS OF TENCH, BREAM AND RUDD.
IBie 21S). IONS Oe EAE SINE
E.AM.& MR. ad nat.del Lith. Anst.v:E_A Funke, Leipzig.
TRYPANOPLASMS OF PIKE ,TENCH, AND BREAM.
TRYPANOSOMES OF TENCH.
JES Se WLIO, AEM WW,
Litn. Anst.v. EA Funke, Leipzig.
H.AM.& MR. ad nat.del
TRYPANOSOMES OF EEL AND PERCH.
1909. } BLOOD-PARASITES OF FRESHWATER FISHES. 33
question of the transmission of these parasites, I thought it worth
while to publish my investigations at the point they have reached,
leaving the subject in a condition in which it can be taken up
again and carried further by myself or by anyone else working in
the same region,
Methods and Technique.
In examining the blood of fishes for the parasites, it 1s not
difficult, when dealing with a fish of fair size, to take the blood
from the gills of the fish without materially injuring it. There
is always a risk, however, that the blood obtained in this way may
contain a certain amount of water mixed with it. To obtain
good smears I found it best to sacrifice the fish. Having killed it
by a smart blow on the head with a blunt instrument (best a
stout rod of hard wood) I opened up the pericardial region and
took the blood from the heart by means of a fine capillary glass
tube, thrust through the wall of the heart; the blood was allowed
to run up into it, and then blown on to the slide or coverslip.
The whole process has to be done quickly, since fish-blood
coagulates very rapidly. Fish which are used in this way for
making blood-smears suffer no detriment in respect to their
culinary properties and need not be wasted.
Much has been written lately about methods of fixation of these
blood-parasites, especially with regard to the procedure most in
vogue, of drying smears of the blood. That the process of drying
affects the minute structural details cannot be doubted and can be
demonstrated easily. It should, however, be pointed out that the
effects of drying, so far as trypanosomes are concerned, differ
greatly according as the drying is done before, or after, fixation
with some histological reagent. Very instructive in this respect
are the species of the genus Zrypanoplasma. ‘Their soft proto-
plasmic bodies become greatly deformed if dried before fixation ;
in this respect they contrast strongly with the species of 7rypano-
soma. In some fishes, for example the pike, tench, and bream,
it is common to find both trypanosomes and trypanoplasms in
the blood; in smears dried before fixation, the trypanosomes
may be found quite satisfactory in form and generai structure,
while the trypanoplasms side by side with them are deformed
almost beyond recognition; compare figs. 25, 26, and 59, show-
ing trypanosomes and trypanoplasms of the pike, from the same
slide; figs. 32, 53, 54, showing similar conditions in the blood
of the bream. In some cases, however, a trypanoplasm may,
apparently, flatten down evenly and thus give a fairly reliable
representation of its natural form and structure (compare figs. 37
and 38, from the same slide); such cases, however, are in my
opinion to be regarded as accidental and exceptional, and the rule
is that trypanoplasms when dried become much deformed. It is
therefore a matter of astonishment to me that Keysselitz should
have relied so much on material dried before fixation for his
1*
4 PROF. E, A. MINCHIN ON PROTOZOAN [Jan. 12,
lengthy and detailed study of the life-history of Trypanoplasma
cyprini (“ borreli”). It is, perhaps, for this reason that Keysselitz
was unable to distinguish more than one species of Trypanoplasma.
The deformation subsequent on drying is entirely avoided if
the smear be placed as quickly as possible, immediately after
smearing and before any appreciable amount of drying has taken
place, into a stoppered tube or bottle containing a few drops of
osmic acid, 4°/, solution. It should be exposed to the vapour for
30-60 seconds, and then transferred, with or without drying, to
absolute alcohol. After the fixation with osmic vapour the drying
does not deform even the trypanoplasms, and the natural form,
size, and appearance of these flagellates are, in my opinion, better
preserved by this method than by any other. The osmic-fixed
smears stain well with Giemsa’s stain, but are apt to stain rather
too darkly and, in the case of large fleshy forms of the parasites,
to become very opaque. Speaking generally, they should be
stained for a much shorter time than the preparations dried off
before fixation.
The weak point of the preparations stained by the Romanowsky
method, whether dried before or after fixation, or not dried at any
time, lies in its effects on the nuclear apparatus. I hope to discuss
this point in greater detail in a memoir which I am preparing on
the structure of Trypanosoma lewisi. I will say here only that
the Romanowsky stain, m its various modifications, gives results
with regard to nuclear structure which are not capable of uniform
interpretation, and which, in my opinion, are untrue and mis-
leading. I believe the defects of the stain to be due principally
to the fact that the red dye or dyes in the stain are precipitated
not only in, but around, certain objects in the preparations ; with
the result that the kinetonucleus, for example, appears many times
its real size, while in the trophonucleus minute, almost ultra-
microscopic granules become enlarged to coarse granules obscuring
the true structure. This effect is not due to the drying, since it
is to be observed in preparations stained by the Romanowsky
method and mounted in Canada balsam without being dried at
any stage of the process. I believe, however, that the process of
drying, whether before or after fixation, may and does add greatly
to the falsity of the nuclear results obtained by the stain. While
osmic vapour fixes very perfectly the cytoplasmic portions of the
body, it apparently leaves the nuclear constituents unfixed and in
a fluid condition. Hence, while the general structure of the body
is very perfectly preserved, even when dried, after osmic fixation,
the nucleus remains in a fluid state and hable to deformation.
That, at least, is the impression which a comparison of different
methods gives me.
For interpretation of the results obtained ‘by the Romanowsky
stain, it is very instructive to compare the totally different results
obtained by staining the same objects with Heidenhain’s iron-
hematoxylin. For this it is necessary, however, that the smears
snould be suitably fixed and that they should have never been
1909. | BLOOD-PARASITES OF FRESHWATER FISHES. 5
dried. JI have tried various fixatives and get the best results with
sublimate-acetic (HgCl, saturated in water 95 volumes, glacial
acetic 5 volumes), Schaudinn’s fluid (HgCl, saturated in water
2 volumes, absolute alcohol 1 volume, with addition of a few
drops of glacial acetic), and Mann’s picrocorrosive-formol solution.
The first two fixatives may be used directly on the wet smears, or
with previous exposure to osmic vapour for a short time. I find
it useless, however, to fix the smears on slides; the reason being
that it is impossible, or at least very difficult, to put them into
the fixatives without making them dive in with one end fore-
most, a process which causes distortion in the wet and still unfixed
elements of the smear. Satisfactory preparations can only be
obtained by these methods of fixation, by making the smear on a
coverslip and dropping it at once with the smear downwards into
the fixative. I hold the coverslip with the fingers of my left
hand, and a glass rod in my right hand ; a drop of blood is placed
by an assistant on the coverslip; I then smear it out immediately
with the glass rod and drop it instantly into the fixative. The
whole process can be done most expeditiously, and with much less
risk of partial drying than when dealing with slides.
In staining trypanosomes with iron-hematoxyln, I find it best
to let the mordant and stain act for a long time. The objects are
first left in the iron-alum solution (34 °/,) over night; they are
then, after brief and rapid washing with distilled water, transferred
to the hematoxylin solution (4°/,) for at least 24 hours. The
whole art of the process lies in the differentiation and extraction
of the stain. My method is to put the coverslip into the iron-
alum until colour is seen to be coming out; then the coverslip is
dipped into tap-water to stop the extraction of the stain, and
examined with moderate magnification (Zeiss Oc. 4, Obj. D). I
the karyosome can be seen sharply and clearly in a trypanosome,
the extraction is sufficient; if not, the coverslip is put into iron-
alum again for a short time. Usually I do several smears of each
sample of blood with different degrees of extraction of the stain.
In most smears there are also thicker and thinner portions, and it
is found that in thicker portions of the smear the stain is not
so quickly extracted from the trypanosomes as in the thinner
portions. Different degrees of extraction of the hematoxylin have
their uses in showing up different points of structure. When the
trypanosomes first come out of the stain they have an even, opaque
black colour. I find that usually the stain is extracted first from
the cytoplasm generally, then from the myonemes ; next from
the coarse granulations of the cytoplasm; next from the flagella
and the blepharoplasts; next from the karyosome, and last of
all from the kinetonucleus which appears to give up the colour
at its periphery first. There are exceptions, however, to this
order, for in some trypanoplasms (e. g. 7’. gurneyorum) the
cytoplasmic granules retain the stain as long as the karyosome.
In one of my preparations which was under-extracted, I found
that the trypanosomes and trypanoplasms showed their form and
6 PROF. E. A. MINCHIN ON PROTOZOAN (Jan. 12,
the Aagella very sharply, but were too opaque for internal struc-
ture; I therefore had the parasites drawn in outline with the
camera lucida, then unmounted the coverslip, extracted more of
the stain, and, after mounting it again, added the details of minute
structure to the drawings already made. ;
With use of iron-hematoxylin in never-dried preparations
suitably fixed, absolutely uniform results are obtained with regard
to the nuclear structure. When two such staining methods as
the Romanowsky stain and the Heidenhain stain give conflicting
results in matters of nuclear detail, no one with any knowledge
of cytological methods would hesitate, I think, to regard as more
reliable the evidence yielded by the Heidenhain stain.
The figures illustrating this memoir were drawn with the camera
lucida to a magnification of 2000 linear, with the exception of the
sketches drawn from living trypanosomes (figs. 77, 94, 95). The
majority of the drawings were executed by my assistant Miss
Rhodes, to whom I desire to express my thanks for her skilled
help in this work,
General Remarks on the Trypanosomes and Trypanoplasms
of Fishes.
It would, perhaps, be more logical if the general account of
these parasites came after the detailed descriptions of the species,
since the conclusions at which I have arrived are founded on the
data which are set forth in the special descriptions. But to many
the general summary of results that is given here will be of greater
interest than the, perhaps, rather wearisome special details, which
therefore I relegate to second place, for purposes of reference for
those specially interested.
From a study of the trypanosomes of fishes in the fresh, living
condition in the blood, I have come to the conclusion that there
are two types of movement: a conclusion which may perhaps be
true of trypanosomes universally. Sometimes they may be seen
to move in a definite direction ; at other times they are seen to be
twisting about in one spot without moving from it. Iterm these
two types of movement travelling and wriggling respectively.
They travel usually with the flagellum forwards ; in the trypano-
some of the eel I have observed progression of this type only.
Some trypanosomes, however, can be seen to travel occasionally
with the flagellum directed backwards ; progression in this manner
appears to me to occur chiefly when the trypanosome is forcing
its way through corpuscles, and never when its path is unobstructed.
Im either case, however, progression appears to be effected chiefly
by flexions of the whole body and by rippling movements of the
undulating membrane, and scarcely at all by means of the free
flagellum.
In wriggling movements the trypanosome simply twists over
and over in S-like curves in one spot, and often appears as if tied
in a knot; this is what I understand Laveran and Mesnil to mean
1909.] BLOOD-PARASITES OF FRESHWATER FISHES. 7
by the expression “ pelotonné.” The object of these movements
is probably simply to increase the animal’s power of absorbing
nutriment &e. from the surrounding medium, by incessantly
changing the surface of contact between the body and the blood-
plasma, just as a Z7’whifex in the mud wriggles incessantly in order
to bring the surface of the body into contact with fresh water and
oxygen. I think it highly probable that when these parasites are
being carried round passively in the circulation they perform
wriggling movements only, having no occasion to travel. On the
slide, however, they are seen sometimes to travel, sometimes to
wriggle ; but when they become moribund they only wriggle.
T'rypanoplasma, so far as I have observed, travels always with
the anterior free flagellum forwards ; [have never seen it go with
the posterior flagellum, that is to say the flagellum which runs
along the edge of the undulating membrane, directed forwards.
If the kinetonuclear extremities of 7rypanosoma and Trypano-
plasma are to be considered homologous, then the direction in
which the former usually travels is the opposite to that im which
the latter invariably progresses. I have not observed in 7rypa-
noplasma anything comparable to the wriggling movements o!
Trypanosoma ; the former genus has the body relatively much
shorter and less flexible. It can be observed frequently, however,
that a Trypanoplasma when travelling in a certain direction will
quite suddenly bend over on itself and travel in a direction more
‘or less the opposite to that which it took formerly.
A vemarkable feature of some trypanosomes and trypanoplasms
of fish is the great disparity in size between different individuals
in the same blood. This point has already been noticed by previous
observers with regard to the trypanosomes of the pike and the
-eel, both of which have been divided, each into two varieties, distin-
guished as var. parva and var. magna respectively. In the five
fish-trypanosomes examined by me, I find the state of things
different in different species. In the trypanosomes of the bream
-and the tench, which are perhaps one and the same species (figs.
27-32), I could not find any variation in size sufficiently well
marked to be characterized as true dimorphism; but my material
-of these two forms is not so abundant as in other cases. In the
trypanosome of the pike (7'. remaki, figs. 20-26) I found the well-
marked dimorphism described by Laveran and Mesnil, and though
the two forms parva and magna each vary slightly in size and
other characters, they were nevertheless easily distinguished, and
no forms could be found transitional between them. In the try-
panosome of the perch (7. perce, figs. 8-14) I found three principal
types—small slender forms, large stout forms, and intermediate
forms. The small forms and the intermediate forms are connected
by transitions (figs. 10, 11), but the stout forms stand rather apart,
owing to the shortness of the free flagellum (figs. 13, 14), Finally in
the eel, I found every possible gradation between the smallest and
the largest forms (figs. 1-7); there is a very great difference in
size between the two extremes, but the absence of a dividing line
8 PROF. E. A. MINCHIN ON PROTOZOAN [Jan. 12,
between them makes it difficult, if not impossible, to distinguish
the varieties parva and magna.
The significance of this dimorphism or polymorphism is not
clear, and must be explained from the life-history. Two possible
explanations present themselves : first, that the difference between
small and large forms is one of growth ‘and development ; secondly,
that itis a manifestation of sexual differences, small male and large:
female forms being differentiated from an indifferent or inter-
mediate form. The trypanosomes of the pike and the perch rather:
favour the sexual hypothesis, but the state of affairs in the eel-
trypanosome strongly suggests stages of growth merely. Certain
facts that I have observed in the perch have awakened in me the
suspicion that these fish-trypanosomes have some form of multi-
plication in the internal organs of the fish, and that fission of the
type familiar in other trypanosomes perhaps only occurs after a
new infection, just as in Zrypanosoma lewisi fission is only found
in the first week or ten days after inoculation. Fission has very
seldom been seen in fish-trypanosomes ; so far as I am aware, it has.
only been seen in 7”. remaki immediately after inoculation into a
pike (Laveran and Mesnil) and in 7. granuloswm in cultures in
vitro (Lebailly, Franca). The absence of fission-stages in the blood
of fish infected naturally is very striking. The subject is one
requiring renewed investigation.
In the trypanoplasms of the tench (figs. 40-43) and pike (figs.
33-37, 56) I have also observed smaller and larger forms, sharply
distinct not only in size but even in nuclear structure; the large
forms are much less common than the smaller ‘ ordinary” forms,
and both types are divisible into two categories by differences in
the nuclear apparatus, especially the kinetonucleus. According
to Keysselitz the large forms are gametes destined to conjugate in
the intestine of the leech; those with larger kinetonuclei are
regarded by him as gametes of male character, while the forms
with smaller kinetonuclei are regarded as female. In one of the
ordinary forms of the trypanoplasm of the pike I have seen the
only example that has come under my notice, of what is apparently
nuclear division (fig. 62).
It is a moot point, how far different species of trypanosomes.
can be distinguished by morphological characters. As I have
already said, the trypanosomes of the five species of fish studied
by me belong, in my opinion, to four species, namely (1) the try-
panosomes of the bream and tench, (2) of the pike, (3) of the
perch, (4) of the eel. Hach of these four putative species can be
easily: distinguished from the other three by its structural cha-
racters, considered as a whole; I need only refer to my figures.
There is, however, always the possibility to be borne in mind that
one and the same species of trypanosome, when: inoculated into
distinct species of fish, might vary in character in response to:
differences in the environment; this is a point on which experi-
mental evidence is needed. The trypanoplasms of the tench,
bream, and pike seem to me also to constitute three well-
1909. | BLOOD-PARASITES OF FRESHWATER FISHES. 9)
characterized morphological species, each quite distinct from the
only specimen of 7. borreli of the rudd that I have seen.
Considering the great variability im size of these parasites, it is
remarkable that so many authors should base their specific descrip-
tions so largely on measurements. A naturalist, for imstance, of
the attainments and experience of my friend M. Brumpt (to whom
I owe my first lesson in the art of staining trypanosomes in
blood-films) can hardly have thought such measurements to be of
any real validity as specific characters.
In considering the minute structure of these parasites we may
begin with the nuclear apparatus. As I have stated above, the
use of iron-hematoxylin as a stain permits of a uniform interpre-
tation of the appearances; and I think it is a very important
point, that it is possible to give in general terms a description of
the nuclear apparatus which will apply to trypanosomes and try-
panoplasms alike. In both cases we find a kinetonucleus and a
trophonucleus, the first named being relatively very large in the
trypanoplasms, small in the trypanosomes. Near the kinetonucleus.
are found the minute blepharoplasts or basal granules of the
flagella, two in 7rypanoplasma, one in Trypanosoma*.
The kinetonucleus in both cases appears as a dense body stained
a uniform deep black after iron-hematoxylin. I have not been
able to detect any structure init. When the stain is over-extracted
it appears to come first out of the periphery of the nucleus and
last of all out of the central part, but this appears to me to be
merely the expression of the dense texture of the kinetonucleus,
and not to indicate any structural difference between central and
peripheral regions. After the Romanowsky stain the kineto-
nucleus appears very much larger than it does after iron-hema-
toxylin; it may be four or five times as large (compare figs. 33—
36 and 57-64; 40-43 and 66-72; 1-7 and 78-85, &.). As I
have stated above, I believe this result to be due to the fact that
the stain forms a deposit round it. With iron-hematoxylin the
size of the kinetonucleus is quite uniform if the stain be not over-
extracted, i which case also the true size is indicated by a clear
space surrounding the central portion stained black.
The blepharoplasts appear usually as very minute dots, scarcely
thicker than the flagellum, after iron-hematoxylin. There
appears to be a band of fibril connecting the kinetonucleus and
the blepharoplasts, which is shown by iron-hzematoxylin, at a
certain degree of extraction (fig. 98); it stains much less deeply
than the kinetonucleus. After the Romanowsky stain the blepharo-
plasts often appear much larger and may be conspicuous ; hence
this stain is useful for demonstrating their existence, although it
* Tadhere, in my descriptions of the structure of these parasites, to the termino-
logy of the organs suggested by me in the ‘Quarterly Journal of Microscopical
Science,’ li. pp. 171-174, text-fig. A; with the difference that I employ the term
“karyosome”’ for the intranuclear body there termed “centrosome.” To judge by
its behaviour during nuclear division, the karyosome of a trypanosome probably
contains a true centrosome or “ centriole.”
10 PROF. E. A. MINCHIN ON PROTOZOAN [Jan. 12,
does not exhibit them in their true proportions. The flagella also
share in the general enlargement after the Romanowsky stain, and
appear very much thicker than they do after iron-hematoxylin,
which shows them as very delicate filaments; they give up the
stain more readily than do the true chromatic structures.
The trophonucleus is very different in its structure from the
kinetonucleus. After iron-hematoxylin (Plates IV. and VY.) it
appears as a clear space, oval or round, limited by a faint but
quite definite membrane, which, when carefully examined, appears
egvanular and uneven in thickness, and is probably composed of
gvanules of chromatin connected together; it is therefore not a
true nuclear membrane, in the sense in which the term is used
for Metazoan nuclei. The trophonuclear membrane is much less
distinct in Zrypanoplasma than in Trypanosoma; a difference
perhaps largely due to its being obscured in the former genus by
the numerous and deeply staining cytoplasmic granulations. In
the space enclosed by the nuclear membrane lie one or more
karyosomes, rounded bodies often very large, and staining very
deeply with iron-hematoxylin. The karyosomes appear to me to |
be simply masses of chromatin; they retain the stain very tena-
ciously. Typically there is a single karyosome placed more or less
centrally, but their size, number, and arrangement vary greatly,
as will be seen by reference to the figures and special descriptions
of the species. In the space between the karyosome or karyosomes
and the nuclear membrane there are found minute chromatin
granules, often disposed so as to leave a clear space round the
karyosome ; the granules themselves are often so minute as to be
scarcely visible.
Where, it may be asked, are the definite chromosomes, eight in
number, surrounding the karyosome, so often described by
Schaudinn, Léger, Keysselitz and others ¢ Where indeed! I can
but describe what I have seen, and I have never seen, after iron-
hematoxylin, more than I have described above. Possibly the
Schaudinnian chromosomes make their appearance at certain
stages of the development or phases of the life-history. I can
only say that they have never revealed themselves to me in any
of my preparations. After the Romanowsky stain, it is true, a
ereat variety of appearances can be seen, impossible to interpret
in a uniform manner : sometimes the whole tr ophonucleus appears
an even red mass, sometimes it shows coarse granulations disposed
in various ways; in all cases its great difference in size, structure,
and appearance from what is seen in the iron-hematoxylin pre-
parations is quite bewildering. I have put forward above, in the
section dealing with technique, what I believe to be the expla-
nation of this. I hope to discuss the whole question much more
fully in dealing with Trypanosoma lewisi elsewhere.
Liihe has given * two figures of the trypanoplasm of the carp
stained with iron-hematoxylin. He figures the trophonucleus as
* In Mense’s ‘ Handbuch der Tropenkrankheiten,’ iii. p. 83, fig. 5.
1909. ] BLOOD-PARASITES OF FRESHWATER FISHES. i
a simple black rounded mass. J am of opinion that what he has
figured is simply the karyosome, and that the peripheral portion
oe the nucleus is not shown. The nuclear space and membrane
are often not at all distinct, as I have said above, and I
have often seen the trophonucleus appearing just as Liihe has
figured it.
In the cytoplasm there are commonly numerous coarse granu-
lations, which stain like chromatin after Romanowsky, but I feel
strong doubts as to their being chromidial in nature. After
iron-hematoxylin I found that the granules gave up the stain
much more readily than the true chromatic structures, in the
trypanosomes ; but in the trypanoplasm of the pike they held the
stain very tenaciously. Their true nature could only be deter-
mined by the development.
In the trypanosomes of the perch (figs. 96, 97) and eel (fig. 84)
I was able to obtain preparations showing ‘the myonemes after
the iron-hematoxylin stain. I have not been able to see myo-
nemes in any trypanoplasm. J am convinced that to see the
myonemes it 18 necessary to get just the right degree of extraction
of the stain; a shade less extra action, and the myonemes are
obscured by the darkness of the cytoplasm; a shade more, and
the myonemes give up the stain. The darkly staining, opaque
cytoplasm of the trypanoplasms probably makes it much more
difficult to render visible the myonemes than in the case of the
trypanosomes. From my preparations the myonemes appear to
be about eight in number in the trypanosome of the eel, but more
than that in the trypanosome of the perch. I have never been
able to make them out with certainty after the Romanowsky
stain.
With regard to the question of the transmission of the trypano-
somes and trypanoplasms of fishes, on which much light has been
thrown by Brumpt and Keysselitz, I have only a few negative
results to record. I found leeches very seldom on the fish caught
by me; only once on a rudd, and once on a perch, a Piscicola was
found attached, On the other hand almost every fish, of what-
ever species, had Argulus attached to it when caught; sometimes
there would be five or six Avguli on one fish. Argulus is stated
in all the memoirs and text-books dealing with it to feed on the
blood of fishes; it is an active swimmer, and readily leaves one
fish and attaches itself to another. It seemed to me therefore
that Argulus was a creature admirably suited by its habits to dis-
seminate blood-parasites, either by the direct mechanical method,
or witha cycle of development. Moreover, Argulus is beautifully
transparent, and everything in its interior can be seen in the
living animal under the microscope ; it is easy to focus its blood-
corpuscles flowing through the heart and circulating in all parts of
its body-cavity, or to see all the contents of its digestive tract,
without injuring the beast in any way. It would be quite an
ideal form in which to study the development and transmission of
hemoflagellates. IJ took numerous Arguli, kept them hungry for
1 PROF. E, A. MINCHIN ON PROTOZOAN [Jan. 12,
a day or two, and then put them into bell-jar aquaria containing
various fishes. Asa rule the Arguli attached themselves at once
to the fish very firmly. After leaving them for varying periods I
removed the Arguli and examined them, first living and uninjured
under the microscope, and afterwards dissected in salt-solution
and examined fresh. Although the fishes on which the Arqula
fed contained trypanosomes or trypanoplasms in all cases, I never
found a flagellate of any sort or description in the Arguli, how-
ever the experiment was varied; and, moreover, I never in any
case found the Arguli to contain blood-corpuscles. It might be
supposed that an animal so transparent as Argulus would show
the effects of a meal of blood by a change of colour visible to the
naked eye; but in no case could a change of colour be discerned
by any optical means whatsoever. I very much doubt, therefore,
if Argulus feeds on blood, or at least on bloed-corpuscles. Very
possibly only fluid plasma or lymph can pass the very minute
terminal aperture of its proboscis.
My experiments with leeches were greatly hampered by the
fact, in the first place, that I can lay claim to no special knowledge
of the Hirudinea, and secondly, that no general monograph of this
group was accessible to me of later date than that of Moquin-
Tandon (1846). It would bea great boon to those studying these
parasites if a more recent monograph or handbook were in existence.
It is not every student of hemoflagellates who is at the same time,
like M. Brumpt, a first-class expert on leeches. I obtained a few
specimens of Piscicola, but found nothing in them of flagellate
nature. I put a Prscicola in an aquarium in company with a
perch ; the leech took no notice of the fish, nor the fish of the
leech. After some days I removed the leech, dissected it, and
found nothmg in it. I also caught in the Broad some other
leeches, which I could not identify accurately, and put them in
with perch ; in all cases the leeches vanished completely, and were
apparently eaten by the fish. Vozld tout!
I can therefore make no positive additions to the transmission-
question, but I hope to return again to this point when opportunity
offers itself.
Description of the Species.
1. TRYPANOsoMA PERCH Brumpt. (Plate I. figs. 8-15; Plate V..
figs. 94-104.)
Brumpt (CR. Soc. Biol. lx. 1906, p. 161) describes this species.
as follows :—‘* Dans le sang de la Perche (Perca fluviatilis). Ce
parasite mesure 57 de longueur totale, dont 16 pour le flagelle,
et 3 de large. Le blépharoplaste est & 1u°5 de l’extrémité posté-
rieure. Le noyau se trouve 4 égale distance de la racine du
flagelle et de la partie postérieure.”
I identify the trypanosomes found by me in the Perch as
7’. perce Brumpt solely on the ground of occurrence, since Brumpt’s.
description might apply to almost any trypanosome, and fails to
1909.] BLOOD-PARASITES OF FRESHWATER FISHES. 13
note any of the peculiarities of the trypanosome of the perch, or
to take into account its variations of size and structure.
Trypanosomes were found more or less abundantly in almost all
the perch examined by me. In a few of these fish none was seen,
or they were found only after more prolonged searching ; as a
rule, however, they were found at once and were present in con-
siderable numbers. If a fish was not well infected, I did not
trouble to make smears of it or to examine it further. Hence
the few cases in which I did not find trypanosomes were probably
cases in which careful search would have revealed the presence of
the parasites in scanty numbers. I doubt if any of the perch
were really entirely free from them. From the perch I examined,
I gained the impression that the trypanosomes were rather more
abundant in small or medium-sized fishes than in those of the
largest size. I did not find trypanoplasms in any of the perch
examined *,
Examination of the blood freshly drawn from the perch showed
that the trypanosomes differed considerably in size. They may
be divided, speaking generally, into large and small forms. The
large forms were much the most abundant; the small forms were
very scarce. The large forms when seen living (figs. 94, 95)
appeared stout and sluggish, as a rule of considerable size, but
showing marked variations in this respect, some being smaller
and more active. They wriggle incessantly but do not travel
much. The body is spindle-shaped; one end, which bears the
flagellum, is greatly attenuated and sharply pointed ; the other
is also drawn out, but is much less attenuated and appears to end
more bluntly. At the blunt end a distinct, very refringent dot
could always be seen, doubtless representing the kinetonucleus ;
it appeared light at a high focus, black at a low focus. Some-
times two small dots could be seen (fig. 94), in which case the
second was probably the blepharoplast. The nucieus could be
seen distinctly as a rounded clear space in which the karyosome
appeared as a darker spot, not very refringent. A short way
behind the nucleus there was seen in some specimens a distinct
dot or grain, much more refringent than the karyosome, and
apparently lodged in a clear space or vacuole (fig. 95); this body
was not always seen. The undulating membrane and flagellum
could be clearly distinguished, especially when the parasites were
moribund and becoming slower in their movements, after being
under observation for some hours.
The trypanosomes show a great tendency in the living state to
twist and roll themselves up, in a way that calls to mind a snake
of stout build, such as a python. Their movements were of two
kinds, which have been distinguished above as wriggling and
travelling movements respectively. When wriggling they simply
twist over and over, throwing themselves into $-like curves,
without changing their place. When travelling the body is
* Keysselitz (Arch. Protist. vii. 1906, pp. 2, 3) records the occurrence of both
‘trypanosomes and trypanoplasms in Perca fluviatilis.
14 PROF. E. A. MINCHIN ON PROTOZOAN [Jan. 12,
stretched out, with the undulating membrane rippling actively,
the undulations commencing at the flagellar extremity and pass-
ing backwards down the body. They travel usually with the
flagellum forwards and the body straightened out; the movement
is effected apparently, chiefly by the undulating membrane; the
blunter posterior prolongation of the body meanwhile performs
curling or wagging movements which appear to be of a passive
kind, the result of the movements of the undulating membrane.
T saw a trypanosome, travelling in this manner on a slide under
a coverslip, approach and go straight through a closely packed
mass of blood-corpuscles, flagellum foremost and body straight.
The same trypanosome was seen also to reverse its movement and
move with flagellum directed backwards; the blunter extremity
of the body, now foremost, performed movements which appeared
to be active in nature, and to assist the body in penetrating
forwards. ‘This was observed also in other cases. ‘Trypanosomes
which were travelling actively were observed to come to a stand-
still and perform simply wriggling movements ; conversely,
trypanosomes which have been wriggling for some time may
start off and begin travelling.
The smaller forms were very similar to the large forms m
their general appearance and the character of their movements,
and only differed in their small size, pronounced slenderness, and
much greater activity and rapidity of movement.
The trypanosome of the perch does not seem to possess the
extraordinary vitality of 7’. granulosum of the eel. 1 found that
the trypanosomes did not live more than a few hours an vitro. I
made an observation, however, relating to their vital powers
which perhaps is not without interest. From a perch which was
very well infected, and from which I made a number of smears
(fig. 11 was taken from this fish) I took three drops of blood;
one drop was simply placed on a clean slide and covered with a
cover-glass; the second drop was placed on a slide and mixed
with a drop of Laveran’s salt-citrate solution, and then covered ;
the third drop was put on a slide and mixed with a drop of tap-
water, and then covered. The three drops were studied from
time to time. In the drop mixed with water the trypanosomes
showed greatly increased activity after about an hour, travelling
with great rapidity in the hemolyzed blood; after four hours
they were still very active, but not quite so active as they had
been earlier. After five hours I slipped off the coverslip, smeared
out the blood, dried it, fixed it with absolute alcohol, and stained
it with Giemsa’s stain. The trypanosomes stained well and
appear perfectly normal. In the drop of blood mixed with salt-
citrate solution the trypanosomes also remained active but some
of them were changed in form after four hours, the body becoming
pear-shaped, with most of its substance aggregated at the hinder
end. In some cases the trypanosomes appeared as if anchored
by the hinder end; the anterior end of the body lashed round in
all directions, but seemed unable to move the inert posterior
1909. | BLOOD-PARASITES OF FRESHWATER FISHES. 15.
mass. After about five hours I slipped off this coverslip also and
made a preparation of the blood, but it only shows trypanosomes
very badly preserved and stained, and apparently quite degenerate
in structure. In the control drop of pure blood the trypanosomes
were all dead or moribund after four hours, and I did not proceed
further with it. If any conclusion can be drawn from these ex-
periments, it is that the trypanosomes live, and remain normal,
longer when tiie blood iv vitro is mixed with water than when it
is mixed with salt-citrate solution or left pure; a result which I
certainly did not expect.
In preparations the trypanosomes of the perch show a con-
tinuous gradation of sizes from the smallest to the largest (figs. 8-
14); the larger forms being, however, by far the commonest.
The best means of classifying them is by the free flagellum, which
in the large stout forms is very short (figs. 13, 14), but in the
medium-sized (fig. 12) and small forms (fig. 8) is much longer.
The cytoplasm stains a very deep blue with the Giemsa stain, so
deeply in fact that it is very difficult to obtain satisfactory pre-
parations of the stout forms; they appear often as bluish opaque
masses in which the intensity of the stain obscures all details of
structure. In the same preparations, on the other hand, more
slender forms may be found perfectly stained. The cytoplasm in
the largest forms usually appears blotchy, with lighter and darker
parts, often with tiny vacuole-like spaces, not very sharply limited.
Any of the forms may contain red-staining granules to a greater
or less extent, sometimes very numerous, sometimes absent alto-
gether. The granules in question are of fair size and more or
less irregular in form. In one specimen I saw them frequently
in pairs, and sometimes rod-shaped, suggesting division (fig. 104) ;
and the idea occurred to me that they might perhaps be intrusive
organisms of the nature of Bacteria. In never-dried preparations.
staloed with iron-hematoxylin the cytoplasm appears more or legs
evenly and coarsely granular, according to the degree of extraction
of the stain (figs. 96-99). ‘The above-mentioned granules are not
brought out by this stain. I see, therefore, no reason for re-
garding them as chromidia.
The nucleus appears, in the smears stained by Giemsa’s method
(figs. 8-14), as a red patch, often obscured and difficult to make
out clearly in the large stout forms. It varies in size with the
dimensions of the trypanosomes, and is much larger in the large
forms. With the Romanowsky stain the details of nuclear
structure appear to vary greatly; sometimes a distinct sharply
limited karyosome, stained a deeper red than the rest of the
nucleus, can be made out (fig. 13), sometimes not. It would
appear as if the method of drying had the frequent result of
distorting or breaking up the fluid or plastic karyosome, thus
producing different appearances in different cases. On the other
hand, in never-dried smears stained with Heidenhain’s iron-
hematoxylin, the structure of the nucleus appears quite uniform
in all its principal features (figs. 97-102), in all cases, and can be
16 PROF. E. A. MINCHIN ON PROTOZOAN (Jan. 12,
described in quite general terms. It is seen that the nucleus is
an ovoid or nearly spherical space limited by a delicate membrane.
In the interior is a deeply staining karyosome, which may be
spherical, ovoid, pear-shaped, or even dumbbell-shaped. The
karyosome is always large, and sometimes so large as to nearly
fill up the entire nuclear cavity and appears to be immediately
surrounded by a clear space. The remainder of the nucleus is
occupied by a faintly granular material, which, owing to the
excentric position of the karyosome, forms usually a crescentic
area on one side of the nucleus. In this area coarser dots of
chromatin can be made out, especially in the neighbourhood of
the nuclear membrane, which is probably composed also of chro-
matin. No details of structure could be seen in the karyosome
itself.
The foregoing statements apply to the nuclear structure of the
large or medium-sized forms, since I was not able, unfortunately,
to find any of the small forms, always rare, in my preparations
stained with 1ron-hematoxylin.
The kinetonucleus appears as a rounded or ovoid mass in
preparations stained by the Romanowsky method. It is larger,
both absolutely and relatively, in the very small trypanosomes
than inthe large. In preparations stained with iron-hematoxylin
it appears either rod-shaped or rounded in form, but in either
case very much smaller in size than it appears when stained by
the Romanowsky method. If the hematoxylin be not much
extracted, the kinetonucleus is often difficult to distinguish from
the blepharoplast, the two together appearing to form a single
mass of triangular or irregular outline. If, on the other hand, the
extraction of the hematoxylin be carried too far, the blepharo-
plast and flagellum become completely decolorized and the
kinetonucleus appears as a very sharply defined and deeply
stained body, im which no details of structure could be dis-
tinguished.
The flagellum arises from a distinct blepharoplast, which can
almost always be seen clearly, and usually lies close beside the
kinetonucleus. In some cases, however, the blepharoplast and
kinetonucleus are separated by a considerable interval, and then
appear to be connected by an ill-defined band of material staining
more lightly (fig. 98). In the living condition also, as stated
above, I observed in some cases two separate grains, which appa-
rently represented the kinetonucleus and blepharoplast.
The flagellum arises direct from the blepharoplast and runs
along the edge of the undulating membrane in the usual way.
As already stated, the free flagellum is short in the large stout
forms, long in those of medium or small size. It is worthy of
note that in the smaller forms the free flagellum ends distally in
a distinct grain or dot. The undulating membrane is shallow,
with many small pleats as a rule, in the stout forms with short
free flagellum (fig. 13); but im all forms with a long free flagellum,
A909. BLOOD-PARASITES OF FRESHWATER FISHES. i
-and especially in the smallest forms, the undulating membrane is
very deep, and stands out far from the body, with fewer and
larger pleats (figs. 8-12). It generally appears clear, but in some
cases the granules of the cytoplasm can be seen extending up into
it, forming a distinct contour-line close under, but quite separate
from, the flagellum (fig. 12).
_ In one of my preparations, which had been fixed first with
osmie vapour and then with Schaudinn’s fluid, without drying,
and stained with iron-hematoxylin, I found the trypanosomes
showing distinctly striations which are doubtless to be explained
as myoneme filaments (figs. 96, 97). They are to be made out
on both surfaces of the body running spirally, and hence appear
to cross in the drawing, but in the object they are seen at
different foci on the two surfaces. In one trypanosome they
appear to run in couples (fig. 96). In another, which was much
bent up and probably contracted, the myonemes can be seen on
one surface running to a very convex edge, where they appear in
optical section as distinct grains, and from this point they can be
traced again on the other surface (fig. 97). The exact number
was difficult to make out; fig. 96 indicates that there are in all 8,
or 4 couples, but in fig. 97 there appear to be more than this.
To these myonemes may be referred the active wriggling move-
ments of the trypanosomes.
In some of my preparations stained with Giemsa’s stain I found
very broad forms of the trypanosome. Jam convinced that these
forms are simply stout forms of the trypanosome deformed and
flattened out by drying. In slides fixed with osmic vapour before
drying I find them only at the edge, or in very thin parts of the
film, that is to say in places where it is difficult to avoid a sight
amount of drying taking place. ‘Two other points are to be noted
in favour of this conclusion; one is that the very broad forms
are not so opaque as the stout forms which do not show any
flattening; the other is that the nucleus is more or less consider-
ably elongated in the transverse direction, having evidently shared
in the increase of breadth produced by the flattening.
In one preparation, fixed wet with osmic vapour and stained
by Giemsa’s method, I found a trypanosome apparently encysted
(fig. 15). No trace of a flagellum was to be seen, but the body is
rounded off and surrounded by an envelope staining a faint
bluish tinge. The shape of the nucleus indicates perhaps that a
slight amount of flattening has taken place. No other similar
stage was found.
In preparations from a perch which showed abundant trypa-
nosomes in its blood, I found, on two separate slides, two bodies
resembling heemogregarines (fig. 16); they were free in the blood-
plasma and strongly resembled the free vermicules of these para-
sites. It is well known that hemogregarines occur commonly in
marine fishes, but in freshwater fishes they have only been found
in the eel. I searched long, but in vain, for intracorpuscular
Proc. Zoon. Soc.—1909, No. IT. 2
18 PROF. E. A. MINCHIN ON PROTOZOAN [Jan. 12,
stages similar to the vermicules; I found, however, in the large
uninucleate * leucocytes, bodies which I Aue at Aret to be para-
sites: rounded bodies staining a faint pink (after Giemsa’s stain),
with a central darker grain or two such grains (figs. 17-19). I
could not find, however, anything in the least transitional to the
vermicule-like bodies, and I do not think now that the bodies in
the leucocytes are of parasitic nature. In the corresponding
leucocytes of the tench (figs. 45-48) there are to be found
commonly, but not invariably, round pink-staining bodies, some-
times a single one of variable size, sometimes two, three, or four
such; they are clearly vacuoles containing some substance which
has stained, probably, with the eosin of the Giemsa stain, and I
think it very probable that the bodies in the leucocytes of the
perch are of a similar nature.
If the vermicules, however, are not stages of a hemogregarine,
in what light are we ‘tor egard them ¢ Since they were found in the
same blood as the encysted trypanosome already described (fig. 15),
the idea occurred to me that perhaps the stout forms of Zrypano-
soma perce might eneyst in the internal organs and undergo
multiplication to form the vermicule-like bodies : these in their
turn might acquire flagella and so give rise to the smallest forms
of the trypanosome, which by growth into the large forms would
complete a cycle of multiplication in the fish. The rarity of
fission-stages of the trypanosomes of fish is remarkable; I have
never seen a fish-trypanosome in division, but Laveran and
Mesnil have described fission of 7’. remaki in two pike infected
artificially, and Lebailly and Franga have, as stated elsewhere,
described fission in 7’. granulosum from cultures in vitro. It is
therefore quite possible that fission may be a process restricted to:
certain parts of the life-cycle, and that the usual mode of multi-
plication in fish-trypanosomes may be such as I have indicated
above. In Zrypanosoma lewisi, for example, fission is only found
during the first week or so after moculation; there is then no.
further multiplication, the trypanosomes being all of one size and
type. On the other hand, tish-trypanosomes usually exhibit
marked variations in size which are very suggestive of growth
from the smallest to the largest forms. I desire to make this
suggestion cautiously, as the data on which it is founded are
obviously quite inadequate to establish it. I may point out,
however, that the blood in which the encysted trypanosome and
the vermicule-like bodies were found, was taken from the heart
of the fish with a capillary glass tube, and it is quite possible that
the tube in passing through the walls of the heart may have
taken up bodies which were not free in the general circulation,
but contained in the wall of the heart itself.
* Commonly termed “mononuclear” ; a barbarous etymological compound. The
adjective “nuclear” means “of or relating to the nucleus”; not “possessing a
nucleus.”
1909. ] BLOOD-PARASITES OF FRESHWATER FISHES. is
2. TRYPANOSOMA GRANULOSUM Lay. & Mesn. (Plate I. figs. 1-7 ;
Plate V. figs. 78-93.)
The trypanosome of the Hel has been seen by many observers,
and there is no other fish-trypanosome which has been the object
of so many memoirs. According to Laveran and Mesnil, the first
description of this parasite was by Sabrazés and Muratet, but the
earlier work of these authors is not accessible to me. The earliest
memoir on the subject with which I am acquainted is that of
Laveran and Mesnil themselves (1902), in which they figure and
describe the parasite and name it 7’. granulosum. Lebailly (1906)
gave a detailed description, with two figures, of this trypanosome
and distinguished two varieties, magna and parva. In the same
year Brumpt described the transmission of the trypanosome of
the eel by the leech Hemiclepsis, and the development that the
parasite goes through in this leech. Finally, Franca (1907) has
devoted a memoir to this trypanosome.
Since so much work has been done on the trypanosome of the
eel, I did not pay much attention to it, not expecting to be able
to add much to our knowledge on this subject. I obtained four
eels caught in the Broad and examined one of them, which I
found to contain trypanosomes most abundantly, more so than
any fish I have yet examined. I made a number of smears and
preparations from this eel and then sent all four eels to the
kitchen, to be prepared for my next meal; it was not found that
infection with trypanosomes impaired appreciably their gastro-
nomic properties. I much regretted afterwards that I did not
examine the other three eels to see if they were equally well
infected.
Sabrazes & Muratet (1904), Lebailly (1906), and Franea (1907),
have all drawn attention to the extraordinary vitality of this
trypanosome, and the fact that if a drop of blood containing them
be sealed up on a slide under a coverslip, the trypanosomes will
live for several days and multiply. I have not imitated these
experiments, but I had a drop of blood under a coverslip, not
sealed up in any way, in which the trypanosomes were moving
actively after 24 hours, less actively after 48 hours, and feebly
after 72 hours.
I spent some time watching the movements of this trypano-
some, which are extraordinarily snake-like. As in the case of
T. perce, the movements are of two kinds, conveniently distin-
guished as wriggling and travelling. They may wriggle for a long
time without changing place to any considerable extent, and then
they may suddenly start travelling. When they become weaker,
after 24 hours under a coverslip, they only wriggle and do not
travel, and as they become moribund the wriggling movements
become weaker and weaker, till on the third day they are very
feeble. When wriggling they twist over and over in S8-like
curves, appearing at the first glance like a writhing knot. When
travelling, I observed them always progressing actively in a
definite direction, flagellum forwards, the body twisting from
Q#
20 PROF, E. A. MINCHIN ON PROTOZOAN [Jan. 12
side to side in even curves like a snake. They frequently stop
suddenly, and wriggle actively in one place like an excited earth-
worm, and then start off again in a new direction. J never saw
them ‘travel with the flagellum directed backwards. The un-
dulating membrane is very distinctly seen, and shows beautiful
rippling movements. Progression appears to me to be effected
chiefly by twists and turns of the body, aided doubtless by the
undulating membrane ; the flagellum has perhaps chiefly a tactile
or euiding function. ‘The kinetonucleus is very distinct in the
ivi condition as a refringent granule a short way from the
hinder end. The trophonucleus is more difficult to see during
life.
Although my preparations were made only with the object of
adding to my collection, they show some points of interest, which
I will state briefly.
In the first place my preparations show trypanosomes of all
sizes from very small to very large (figs. 1-7). The contrast
between the two extremes of the series is surprisingly great, but
all possible transitions from one extreme to the other are to be
met with. Nor can I find any definite structural features to
separate sharply the large and small forms, as can often be done
in trypanosomes of other species. The large forms are very
granular, while in the smallest granules are absent or scarce, but
this feature shows both variations and transitions; the large
forms have more pleats in the undulating membrane than the
smaller, but the pleats are about the same depth and simply
increase in number with the length of the trypanosome. The
free flagellum bears about the same proportion, as regards length,
to the rest of the body in both large and small fo'ms, though im
some large forms it is perhaps relatively shorter. The differences
in the nucleus, to which I shall refer presently, are also differences
due to a gradually increasing complexity of structure. I cannot
therefore find characters to separate sharply the varieties magna
and parva of Lebailly.
A very marked feature of this species, when stained by the
Romanowsky method, is the occurrence of numerous granules in
the cytoplasm, a peculiarity to which it owes its name. The
granules stain purple or blue with Giemsa’s stain. In prepara-
tions stained with iron- -hematoxylin, the granules are only to be
seen when the stain is not sufficiently extracted (fig. 83); when
the stain is most satisfactory and the flagellum and nuclei are
sharp and distinct, the characteristic granulations are not to be
seen. One of my coverslip-smears is rather uneven and thicker
in some parts than in others. In the thicker portions the stain
is less extracted than in the thinner parts of the smear, as
commonly happens, and hence different degrees of extraction can
be found in the preparation. Only a very few trypanosomes show
the granules stained and standing out sharply; with too slight
extraction of the stain the granulations are obscured by the
darkness of the cytoplasm. If the stain 1s very much over-
'
1909. } BLOOD-PARASITES OF FRESHWATER FISHES. 21
extracted, as in one of my preparations, the colour comes out of
everything except the kinetonucleus; when the extraction has
not gone quite so far, the karyosome of the trophonucleus also
retains the black colour. The granulations of the cytoplasm are
not, in my opinion, to be regarded as chromidia, since they give
up the stain very readily, while the true chromatic structures
retain it very tenaciously.
Franga has drawn attention to the peculiar structure of the
nucleus in the variety magna of this trypanosome. He writes:
“hes moval, ses. « s'éloigne par sa structure de celui de presque
tous les Trypanosomes. Au sein d’une substance incolore on voit
la chromatine formant d’ordinaire deux parties bien distincteés ; un
arama OLOCE eae. dun rouge vif, situé dans la partie du noyau
la plus rapprochée du blepharoplaste; et un grand nombre de
granulations petites et réguliéres, ou un reticulum chromatique,
vers le coté du flagelle. Dans quelques parasites, plus rares, on
ne voit pas le grand bloc de chromatine et alors il existe dans
la substance nucléaire incolore une série de granulations dis-
posées de fagon 4 former un cordon plus au moins sinueux.” “ Au
contraire de ce qui existe dans le noyau des Trypanosomes de la
var. magna, dans ceux de la var. parva il n’y a pas une distinction
nette entre les portions chromatique et achromatique.”
Franga’s observations appear to have been made on trypanosomes
stained by the Romanowsky method, and my preparations coloured
with Giemsa’s stain confirm his statements, But I do not believe
it is possible to get a coherent or intelligible idea of the structure
of the nucleus with this method of staining. My preparations
stained with iron-hematoxylin show appearances quite different
from those seen after use of the Romanowsky stain, and at the
same time permit of a uniform interpretation of the structural
details. The nucleus appears very much smaller and more
compact after iron-hematoxylin than after the Romanowsky
stain; and while it is possible that there is a certain amount of
shrinkage by the former method, [ think there is certainly a
considerable amount of artificial expansion and deformation con-
sequent on the process of drying by the ordinary method of
applying the Romanowsky stain.
In never-dried preparations stained with iron-hematoxylin, the
smallest trypanosomes (figs. 78, 79) show the nucleus as a clear
oval space with a distinct limiting membrane, containing
sharply defined karyosome of elongate-oval form. The karyosome
is far from filiing up the entire nuclear space, which appears clear
or shows very minute granulations. A nucleus of this type of
structure may be found even in trypanosomes of large size
(fig. 87), but the karyosome in such specimens is considerably
larger and fills up nearly the whole nuclear cavity. Asa rule,
however, in trypanosomes only slightly larger than the smallest
that can be found (figs. 80, 81), the karyosome is seen to have
budded off from one extremity, usually from that furthest from
the kinetonucleus, a smaller part. This is the type of nucleus
22 PROF. E. A, MINCHIN ON PROTOZOAN [Jan. 12,
most commonly found, one in which there are two karyosomes,
either subequal, or markedly unequal, in size (figs. 85, 86, 93).
But in many of the large forms the karyosome may be further
broken up into three or even four parts, usually differing greatly
in size (figs. 89-92). Thus, with endless variations in detail, a
similar type of nuclear structure occurs with monotonous regu-
larity in all the preparations stained with iron-hematoxylin ;
always an oval space containing one or more deeply staimed karyo-
somes. What may be the significance of the disruption of the
karyosome I am unable to say.
The kinetonucleus appears after iron-hematoxylin as a rounded
or rod-shaped body of minute size. After staining by the Roma-
nowsky method it appears very much larger and of various shapes
which are seen in my figures. Near it a distinct blepharoplast
can be made out, from which the flagellum arises directly. The
free flagellum is usually of considerable length. The undulating
membrane does not stand out so boldly from the body as in some
species of fish-trypanosomes. In one preparation I saw distinctly
a line under the flagellum (fig. 82), indicating doubtless the limit
of the extension of the endoplasm into the undulating membrane.
Only in one specimen was I able to make out myonemes (fig. 84).
They appeared as delicate lines, about three being visible on the
side of the body, a number which may correspond to the total of
eight alleged to be typical of trypanosomes.
3. TRYPANOSOMA REMAKI Lav. & Mesn. (Plate II. figs. 20-26.)
The trypanosome of the Pike has been described by Laveran and
Mesnil (Arch. Protist. i. 1902, pp. 482-486, figs. 1-9). I have
very little to add to their description of this parasite, which I
found in four out of the five pike examined by me at Sutton
Broad. Laveran and Mesnil have noted that this parasite has a
wide distribution in Europe, its occurrence having been noted by
several naturalists.
Laveran and Mesnil noted two forms which they distinguished
as var. parva and var. magna. 1 found these varieties very dis-
tinctly marked in the pike examined, and without any transi-
tional forms connecting the two extreme types. Both var. parva
and var. magna are subject to slight variations in size, but I found
no difficulty whatever in distinguishing the two types in my
preparations, whether fixed by osmic vapour or by the ordinary
method of drying. The var. parva (figs. 20-22, 25) is clearer,
with but few coarse granulations, its undulating membrane is
shallow and does not stand out sharply from the body, and its
free flagellum is longer, relatively and absolutely, than that of
var. magna (figs. 23, 24, 26), which contrasts with it not only in
greater dimensions, but also in its opaque cytoplasm full of coarse
granules staining reddish with the Giemsa stain, and in its very
prominent undulating membrane. The kinetonucleus also seems
to be slightly larger in var. parva than in var. magna; but I am
not able, unfortunately, to make precise statements concerning
1909. | BLOOD-PARASITES OF FRESHWATER FISHES. 23
the nuclear structure, since in the smears which I stained with
iron-hematoxylin I was unable to find any trypanosomes, though
Trypanoplasma gurneyorum was abundant.
4. TRYPANOSOMA TInca Lav. & Mesn. (Plate II. figs. 27-29 ;
Plate IV. figs. 74-76).
The trypanosome of the Tench was described and named by
Laveran and Mesnil (Trypanosomes et Trypanosomiases, Paris,
1904, pp. 387, 388, fig. li. 2 & 3). Its occurrence had been
noted previously by Doflein (Die Protozoen als Parasiten etc.,
Jena, 1904, p. 71), and since then by Keysselitz (Arch. Protist.
vil. 1906, p. 3). Doflein considered that the parasite seen by him
might be identical with Trypanosoma carassii from Carassius
vulgaris, but Laveran and Mesnil regarded it as a distinct and
new species, and described it as follows :—
* La longueur est de 35 4 en moyenne, la largeur de 24 4a 3
L’extrémité postérieure est conique, peu effilée. Le centrosome,
assez gros, est voisin de l’extrémité postérieure. Le noyau est situé
vers la partie moyenne du corps du parasite. La membrane on-
dulante est large, bien plissée. La partie libre du flagelle est
assez longue.”
I found trypanosomes in all tench examined by me at Sutton
Broad, occurring together with Zrypanoplasma keysselitzi, and
have observed nothing to prevent me identifying them with
Trypanosoma tince of Laveran and Mesnil. This trypanosome
appears to be very uniform in size and structure, so far as my
observations extend ; the two specimens figured by Laveran and
Mesnil differ somewhat in size. In my never-dried preparations
fixed with Schaudinn’s fluid, with or without previous fixation with
osmic vapour, and stained with iron-hematoxylin (figs. 74-76),
the trypanosomes appear constantly smaller than they do in pre-
parations fixed wet with osmic vapour, or dried before fixation,
and stained with Giemsa’s stain; but I attribute this difference to
the effects of the method of fixation.
In osmic-fixed preparations the trypanosome shows, as Laveran
and Mesnil have stated, a very broad undulating membrane,
standing out well from the body, and continued into a free
flagellum of moderate length (figs. 28, 29, 36); the cytoplasm is
opaque in such preparations and usually shows numerous coarse
granules, which stain reddish with Giemsa’s stain, and by the
iron-hematoxylin method appear black and hold the stain fast.
The kinetonucleus appears large after Giemsa; but very much
smaller after iron-hematoxylin, and by the latter method shows
two types of form, rounded and rod-shaped (figs. 74-76). The
blepharoplast is not easily seen in the Giemsa preparations,
owing to the opacity of the cytoplasm, but in iron-hematoxylin
preparations it appears as a sharp dot near the kinetonucleus.
The trophonucleus appears after Giemsa stain as an oval patch
showing no details of structure ; after iron-hematoxylin it appears
as a rounded or oval space, not very sharply limited, containing
a4 PROF. E. A. MINCHIN ON PROTOZOAN [Jan. 12,,
a very large round karyosome. In one preparation the karyosome
appeared to fill the whole nuclear space and to be of irregular
form (fig. 75); a result perhaps due to insufficient extraction of
the stain.
5. TRYPANOSOMA ABRAMIS Lay. & Mesn, (Plate IT. figs. 30--32.)
The occurrence of trypanosomes in a Bream was noted by
Laveran and Mesnil in 1902 (Arch. Protist. i. p. 478) and the
species was named by these authors, but without any figure or
description, in 1904 (Trypanosomes et Trypanosomiases, p. 388).
Keysselitz also noted the occurrence of this parasite (Arch.
Protist. vii. p. 3). According to Brumpt, the trypanosome of
the bream goes through its development in the leech Hemiclepsis.
In my preparations of the blood of the bream from Sutton
Broad I have found a trypanosome very sparingly; three
Specimens altogether, one from one fish, two from the other.
The trypanosome is of large size and presents no character by
which I can distinguish it morphologically from Zrypanosoma
tince. 1 consider it very probably identical with this species ;
in which case the name ftincw has priority over abramis by one
page; but in view of the scantiness of my observations I refrain
from taking the step of merging the two species into one.
6. TRYPANOPLASMA GURNEYORUM, sp. n. (Plate IT. figs. 33-39 5.
Plate LV. figs. 56-65.)
The occurrence of a trypanoplasm in the Pike has only been
noted, so far as T am aware, by myself (Quart. Journ. Mier. Sei,
lii. p. 253, text-fig, B). I found it abundantly in all pike
examined at Sutton Broad, usually occurring together with
Trypanosoma remaki Lay. & Mesn.; in one pike, however, I
found a very abundant infection with the trypanoplasm, but no
trypanosomes. I propose to name this trypanoplasm 7’. gurney-
orum, in honour of Messrs. Eustace and Robert Gurney, in whose
laboratory this work was done.
Trypanoplasma gurneyorum occurs under two forms, which
I will distinguish as “ordinary ” and “large.” The large forms
appear to be very scarce; I have seen only two specimens so
far, both in preparations from the fish mentioned above, in which
only trypanoplasms were found. The ordinary form of 7”. gurney-
orum (figs. 33-37) is characterized by a fairly broad undulating
membrane and by the shortness of the free flagella. The anterior
free flagellum is searcely more than half the length of the body.
The posterior free flagellum projects only a very short distance
beyond the hinder end of the body ; in some specimens it is rather
longer, but it is always short by comparison with other trypano-
plasms. The trophonucleus is near the middle of the body,
sometimes even in the posterior half. The form of the anterior
extremity of the body shows variations, which are perhaps due
1909.) BLOOD-PARASITES OF FRESHWATER WISHES, 25)
partly to the animal’s movements, partly to deformations induced
by fixation. Sometimes the anterior end appears pointed and the
marginal flagellum runs backwards from its point of origin (fig. 36).
On the other hand, the anterior end may bevery blunt and rounded,
and the marginal flagellum rons first forwards and then back
wards from its point of origin (fig. 33). ‘The second type is found
in an extreme form in preparations dried before fixation (fig. 37),
and is then undoubtedly largely a case of deformation ; the chief
mass of the body seems to flow forwards when dried, with the
result of bringing the kinetonucleus and trophonucle us close
together. In extreme cases the try panoplasin becomes a shapeless
mass, in which the orientation of the bocy is difficult to make out
(figs. 388, 39). It is interesting to note that, Mihi n ycke
defor med in this way beyond recognition oceur in preparations
in which the trypanosomes have their body fiir aad structural
characters preserved quite perfectly.
The large form of Vrypanoplasma gqurneyorum (fig. 56) also
has the free flagella velatively short. The bocy el ae is very
dense and opaque, staining a very deep blue with Gie msa’s stain,
so that it is very difficult to make out the nucleus; in the speci-
men drawn in outline in fig. 56 I could see the kinetonucleus
plainly, especially with green light, but the trophonucleus was
very difficult to distinguish from the cytoplasm, and I am by no
means certain that IT have drawn it correctly.
The most remarkable feature of Trypanoplasma gurneyorwn is
the presence of deeply staining granules in the cytoplasm, They
oceur chiefly towards the hinder end of the body, but are found
also, though more sparingly, up to the anterior extremity. By
the Romanowsky method (figs. 33-39) they stain deeply in a
colour approaching the tint of the nuclei, and more especially that
of the kinetonucleus. By the ivon- hwmatoxylin method they hold
the stain very fast (figs. 57-65), quite as fast as do the nuclei, a
point in which they contrast with the eytoplasinic granules of
Trypanosoma granulosum and 7. perew, he granules are
especially sharp and clear in preparations fixed with sublimate-
acetic and stained with iron-hematoxylin (figs. 57-61), They
are rounded and vary in size from small dots to coarse grains.
Sometimes there are only a few granules, but as a vule they are
very numerous and give this trypanoplasm a very characteristic
appearance,
Ay ie nuclei of Trypanoplasma quimeyorum appear very different
according to the stain used, Romanowsky or iron-heimatoxylin.
The trophonucleus is lodged in ov near the edge of the undulating
membrane, and hence is difficult to make out clearly if the
undulating membrane in this part be folded over the body, It
appears after the Romanowsky stain as a lightly stained patch,
more or less oval in form, showing no detail or only a few
ivvegular granulations, After Won -hematoxylin (figs. 57-64) it
appears as a clear oval space, not very well defined, containing
26 PROF. E. A. MINCHIN ON PROTOZOAN [Jan. 12,
usually a single karyosome. In the ordinary forms the karyosome
is small, often no larger than one of the coarser cytoplasmic
granules, with which it may be confused very easily, so that at
first sight the trypanoplasms appear to have no trophonucleus.
Tn the only specimen of the large forms which I have seen stained
by this method, the karyosome was very large and almost filled
the nuclear space (fig. 65). In one specimen of the ordinary
forms I found what appeared to be nuclear division (fig. 62); two
karyosomes connected bya distinct curved black line, one of them
in the ordinary position of the trophonucleus, the other further
forwards, on a level with the kinetonucleus. In another specimen
the karyosome appeared to be budding off a smaller part (fig. 59),
perhaps as a preliminary to division.
The difference between the nucleus in specimens stained by the
Romanowsky and the iron-hematoxylin methods is very striking.
One sample, however, of Giemsa’s stain which I used gave
different results from all others, for some unexplained reason ;
with this sample the flagellum stained blue instead of red, and
the trophonucleus appeared as a small granuie (fig. 36), just as in
the iron-hematoxylin preparations; the kinetonucleus appeared
very large. Hence all the trypanoplasms stained with this
sample of stain gave the impression, at first sight, of lacking a
trophonucleus.
The kinetonucleus appears very large after the Romanowsky
stain, much smaller after iron-hematoxylin, but in both cases
larger, as a rule, than the trophonucleus. There are two types
of form exhibited by the kinetonucleus : a more rounded or oval,
broader type (figs. 57, 59), and a more elongated, band-hke type
(figs. 61, 64); the latter often appears to have a small piece at
the anterior end constricted off from the main body (figs. 58, 64;
compare Keysselitz, Arch. Protist. vil. p. 36, fig. 45). The
kinetonucleus stains an even black in iron-hematoxylin, a deep
purple after Giemsa, and shows no structural detail. Having
only seen two specimens of the large form of this trypanoplasm,
T am unable to say if distinct types of kinetonucleus are exhibited
by it, as by the large forms of 7. keysselitzi. In front of the
kinetonucleus are two minute blepharoplasts from which the
flagella arise. After the Romanowsky stain the blepharoplasts
appear a fair size and are distinctly seen ; after iron-hematoxylin,
on the contrary, they are very minute and often appear as a single
granule, being close together and difficult to resolve imto two
distinct dots. When the anterior end of the body is drawn out,
the blepharoplasts appear sometimes one behind the other, and
with a considerable interval between them.
The following is an attempt at a brief diagnosis of 7rypano-
plasma gurneyorum:—Oceurs under two forms, ordinaryand large ;
trophonucleus near middle of body, with single karyosome ;
kinetonucleus compact or drawn out, large; anterior flagellum of
moderate length ; free portion of posterior flagellum very short.
Host, Hsox lucius, Norfolk.
1909. | BLOOD-PARASITES OF FRESHWATER FISHES. 27
7. TRYPANOPLASMA KEYSSELITZI, sp. n. (Plate III. figs. 40-44;
Plate IV. figs. 66-73.)
The occurrence of a trypanoplasm in the Tench has been noted by
Keysselitz (Arch. Protist. vii. p. 3), but the species has not been
described in detail, so far as I am aware, or named. Since it
appears to me to be a well-marked species characterised by several
striking peculiarities of structure, I propose to name the trypano-
plasm of the Tench Zrypanoplasma keysselitzi, in honour of
Dr. Keysselitz, who was the first, I believe, to observe it. From
the figure given, however, by Lithe (Mense’s Handbuch, 111. fig. 5,
p. 85) of the trypanoplasm of the Carp, I think it possible that
T’. keysselitzi may prove to be a synonym of 7’. cyprini Plehn.
Trypanoplasma keysselitzi was found im all the tench examined
by me at Sutton Broad, always in company with 7rypanosoma
tince. In some of the tench examined it was very abundant. In
one case I saw as many as four or five in a field (Zeiss Oc.
Obj. D) in a drop of fresh blood under the coverslip. In other
cases the parasites were scarcer, but in no case was I unable to
find them.
The trypanoplasms occur under two distinct forms : one smaller,
which, as it is the most abundant, I will call the ordinary form
(figs. 42, 43); the other, less common, I will refer to simply as
the lar oe form (figs. 40, 41). The two forms are easily dis-
tinguished in the living state in the fresh blood. In preparations
they are seen to be distinguished not only by differences in size
but also by points of structure. The posterior flagellum is con-
tinued beyond the undulating membrane as a relatively very long
free flagellum in the ordinary forms, but the free portion is quite
short in the large forms. There are also differences in the nuclear
structure, presently to be described, between the ordinary and the
large forms, and by this feature the large forms can also be
separated into two types.
The cytoplasm is finely and densely granular, and in the large
form opaque and deeply-staining, but it is relatively free from
the coarse granules which are such a feature of 7. gurneyorum.
In iron-hematoxylin preparations only a few coarse granules are
to be seen, sometimes none at all.
The two nuclei of this trypanoplasm are remarkable, in the
first place, for being situated very close together. The tropho-
nucleus, always placed at the edge of the undulating membrane,
is never more than a short way behind the kinetonucleus, which
it usually overlaps at the hinder end, and sometimes even the
trophonucleus is opposite the middle of the kinetonucleus. The
arrangement of the two nuclei gives this trypanoplasm a very
characteristic figure and appearance.
The trophonucleus shows well-marked variations of structure.
In the ordinary forms, when stained with iron-hematoxylin, it is
2 small oval clear space which contains two distinct karyosomes
(figs. 70-72). These bodies may appear as small dots, looking
like two granules of the cytoplasm, or they may be larger, but
28 PROF. E, A. MINCHIN ON PROTOZOAN [Jan. 12,.
the number is constant, so far as my observations extend. One
karyosome is often slightly larger than the other. When the
trypanoplasm lies in a certain position, the karyosomes may be
quite hidden under the opaque kinetonucleus, and then the tro-
phonucleus cannot be made out at all. In the large forms, on the
other hand, the trophonucleus shows either one large karyosome
which almost fills the nuclear cavity (fig. 69), or one large and
one or two smaller karyosomes (figs. 67, 68). There are further
differences to be observed in the trophonuclei of the large forms
which appear to be correlated with differences in the kinetonuclei.
Usually a large trophonucleus with a very large karyosome is
correlated with a relatively short kinetonucleus (fig. 69). On the
other hand, a small trophonucleus may be found combined with a
very long kinetonucleus (fig. 67). According to Keysselitz the
large forms of the trypanoplasm are gametes, those with the
large kinetonucleus being male forms, and those with smaller
being female. I am not in a position to criticise or comment
upon this statement.
The kinetonucleus is situated on the side of the body furthest
from the undulating membrane, and is remarkable for its great
length, being narrow and band-like, never rounded, oval or pear-
shaped. As already stated, its length varies in different specimens.
With iron-hematoxylin it always stains an even black and shows
no structure: in one specimen which had been very much ex-
tracted (fig. 72) it appeared very narrow and linear in form, with
a clear space in the cytoplasm on one side.
In front of the kinetonucieus are situated the two minute
blepharoplasts from which the flagella arise. I believe them to
be always two im number, but in iron-hematoxylin preparations
they are so minute and often so close together that it is impossible
to resolve them as two granules and they may appear as a single
dot. There appears to be a streak or fibre connecting the
kinetonucleus with the blepharoplasts, but the length of the
connection, that is the distance between the two structures, is
variable.
The following is an attempt at a brief diagnosis of 7rypano-
plasma keysselitzi :—Oceurs under two forms, distinguished by
size and by structural characters; in both forms the two nuclei
very close together at the anterior end of the body; kineto-
nucleus very elongated, trophonucleus small; posterior free
flagellum long in the smaller forms, short in the larger forms.
Host, Tinca vulgaris, Norfolk.
8. TRYPANOPLASMA ABRAMIDIS Brumpt. (Plate ITI. figs. 49-54 ;
Plate 1V. fig. 77.)
The trypanoplasm of the Bream has been described by Brumpt
(CR. Soe. Biol. lx. 1906, p. 164) in the following words :—
* Dans le sang de la Bréme. Vu seulement a l'état frais. Le
corps posséde alors 30u de long sans compter les flagelles, Pantérieur
ayant environ 15, de long, le postérieur 5 et 64 seulement. Ce
1909.] BLOOD-PARASITES OF FRESHWATER FISHES. 29
parasite évolue exclusivement chez | Hemiclepsis et ne passe
jamais dans la gaine de la trompe, ce qui permet de le distinguer
des especes déja connues de la Carpe et des Loches.”
The occurrence of trypanoplasms in the bream has also been
noted by Keysselitz (Arch. Protist. vii. 1906, p. 3).
1 found trypanoplasms fairly abundantly in all the bream
examined by me at Sutton Broad. It is unnecessary to give a
detailed description ; my figures show their general appearance.
The body is rather slender and not very granular. The anterior
flagellum is fairly long, the posterior free flagellum moderately so.
The trophonucleus is about on a level with the hinder end of the
kinetonucleus. I did not staim any preparations with iron-
hematoxylin, so can say nothing about the minute structure of
the nuclei. I did not find any trypanoplasms of large size, as in
other species.
In its characters 7rypanoplasma abramidis appears to be more
or less intermediate between 7’. keysselitzi and 7. gurneyorum, but
distinct from either.
TRYPANOPLASMA BORRELI Lav. & Mesn. (Plate IIT. fig. 55.)
The trypanoplasm of the Rudd (Leuciseus erythrophthalmus) is
the type-species of the genus 7rypanoplasma; it was described
under the name 7’. borreli by Laveran & Mesnil in 1902 (Arch.
Protist. 1. p. 489). The Rudd and Roach are very abundant: in
Sutton Broad, perhaps the two commonest species of fish there, and
Texamined a great number of specimens of each species for blood-
parasites ; only in one rudd, however, was I successful in finding
them. J saw one trypanoplasm in the fresh blood, and made
several smears from the same fish. After very prolonged
searching of the smears a single trypanoplasm was found, which
both from its occurrence in the Rudd and from its resemblance
to the figures of Laveran & Mesnil and Léger, I have no
hesitation in identifying as 7’. borreli. From the single specimen
before me, 7’. borreli appears to resemble 7’. keysselitzi in the length
of its free flagella, but differs in having the trophonucleus further
back (fig. 55).
EXPLANATION OF PLATES I.-YV.'
All figures are drawn with the camera lucida to a magnification of 2000 linear;
with the exception of fig. 77, Plate IV., and figs. 94, 95, Plate V., which are drawn
from the living object freehand.
Prate I,
Figs. 1-7. Trypanosoma granulosum of the Eel, a series to show the gradations in
size between the smallest and the largest forms. All from the same
blood, and all, except fig. 7, from the same slide. Preparations dried
off, then fixed with absolute alcohol and stained with Giemsa’s stain.
Figs. 8-14. Trypanosoma perce of the Perch, various forms. 8-11, small forms ;
12, medium-sized form with long free flagellum; 13, 14, stout forms
with short free flagellum; 8 & 12-14 from preparations dried off
and fixed with absolute alcohol; figs. 9-11, from preparations fixed
with osmic vapour followed by absolute alcohol; all stained with
Giemsa’s stain.
30 PROTOZOAN BLOOD-PARASITES OF FRESHWATER FISHES. [Jan. 12,
Fie. 15. Encysted form of Trypanosoma perce, from the same slide as 9 and 10.
Osmie vapour, absolute alcohol, Giemsa.
Fig. 16. Young form of 7. perce? or vermicule of a hemogregarine? From
the same slide as the last.
Figs. 17-19. Leucocytes of the Perch, showing peculiar enclosures. From a
preparation dried off, fixed with absolute alcohol, and stained with
Giemsa’s stain.
eNom) JUG
Figs. 20-26. Trypanosoma remaki of the Pike. 20-22 & 26, small forms (var.
parva) ; 28, 24, & 26, large forms (var. magna) ; 20-24, from pre-
parations fixed with osmic vapour followed by absolute alcohol; 25, 26,
from preparations dried off, fixed with absolute alcohol, all stained
with Giemsa’s stain.
Figs. 27-29. Trypanosoma tince, from the Tench. 27, dried, absolute alcohol,
Giemsa ; 28, 29, osmic vapour, absolute alcohol, Giemsa.
Figs. 30-32. Trypanosoma abramis, from the Bream. 30, 31, osmic vapour,
absolute alcohol, Giemsa; 32, dried off, absolute alcohol, Giemsa.
Figs. 33-39. Trypanoplasma gurneyorum, from the Pike. 83-36, osmic vapour,
absolute alcohol, Giemsa; 37-39, dried off, absolute alcohol, Giemsa.
Prave IIT.
Figs. 40-44. Trypanoplasma keysselitzi, from the Tench. 40, 41, large forms ; 42—44.,
ordinary forms; 40-43, osmic vapour, absolute alcohol, Giemsa ;
44, dried off, absolute alcohol, Giemsa.
Figs. 45-48. Large uninucleate leucocytes of the Tench, showing peculiar pink-
staining enclosures; preparation dried off, fixed with absolute alcohol,
stained with Giemsa’s stain.
Figs. 49-54. Trypanoplasma abramidis, from the Bream, 49, fixed with sublimate-
acetic (95:5); 50-52, fixed with osmic vapour followed by absolute
alcohol; 53, 54, dried off, fixed with absolute alcohol; all stained
with Giemsa’s stain.
Fig. 55. Trypanoplasma borreli, from the Rudd. Osmie vapour, absolute
alcohol, Giemsa.
PrarE TV.
Figs. 56-65. Trypanoplasma gurneyorum, trom the Pike. 56, large form, from a pre-
paration stained with Giemsa’s stain, all the others from preparations
stained with iron-hematoxylin; 56, osmic vapour, absolute alcohol ;.
57-61 & 63, sublimate-acetic (95:5); 62, 64, 65, Mamnn’s picro-
corrosive with formol.
Figs. 66-73. Trypanoplasma keysselitzi, from the Tench; all from preparations
stained with iron-hematoxylin. 66-69, large forms; 70-73, ordinary
forms; 66, Schaudinn’s fluid; 67 & 70, sublimate-acetic (95:5) .
68, 69, & 71-73, Mann’s picro-corrosive with formol.
Figs. 74-76. Trypanosoma tince, from the Tench, all from preparations stained
with iron-hematoxylin ; 74, Schaudinn’s fluid direct ; 75-76, Schau-
dinn’s fluid preceded by exposure to osmic vapour.
Fig. 77. Trypanoplasma abramidis, trom the Bream, sketched living.
Prare V.
Figs. 78-93. Trypanosoma granulosum, trom the Kel, all from preparations stained
with iron-hematoxylin. 78-82 & 85-93, fixed with Mann’s picro-
corrosive-formol mixture; 83, 84, fixed with sublimate-acetic (95 : 5).
78-80. Small forms.
81-82. Medium-sized forms; in 82 only the posterior part figured, to
show the double line in the undulating membrane, the outer line
representing the flagellum, the inner line representing the limit of the
extension of the endoplasm imto the membrane.
83. Large form, anterior half, the stain but slightly extracted, showing
the cytoplasmic granules ; whole length of the flagellum not drawn.
84. Large forms, anterior half, stain under-extracted, showing,\ the
myonemes ; whole length of the flagellum not drawn.
85. Large form.
86-93. T'rophonuclei of different specimens, all from the same prepar-
ation, and all from large forms.
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1909.] CoPEPODA OF THE THIRD TANGANYIKA EXPEDITION. 3k
Figs. 94-104. Trypanosoma perce, from the Perch, all from preparations stained
with iron-hematoxylin, except 94, 95, 103, 104. 97, 98, fixed with
Schaudinn’s fluid after previous exposure to osmic vapour ; 98-102,
fixed with Mann’s picro-corrosive-formol ; 103, osmic vapour followed.
by absolute alcohol, 104, dried off and fixed with absolute alcohol,
both stained with Giemsa’s stain.
94, 95. Sketch of Trypanosoma perce, large forms, drawn living ;
94, a Specimen showing two dots at the hinder end (kinetonucleus
and blepharoplast ?); 95, a specimen showing a refringent dot,
lodged in a vacuole, close behind the trophonucleus.
96. Specimen showing myonemes on both surfaces of the body, from a
preparation in which the stain was but slightly extracted.
97. Middle portion of another specimen, very contracted, showing
myonemes, from the same preparation as the last.
98. Specimen showing a large blepharoplast, connected with the kineto-
nucleus (or division of the kinetenucleus 2).
99. Another specimen from the same preparation as the last.
100-102. Trophonuclei of different specimens.
103, 104. Stout forms drawn to show the distribution of the cytoplasmic
eranules ; in 104 many of the granules are in pairs, as if dividing or
recently divided.
2. Zoological Results of the Third Tanganyika Expedition,
conducted by Dr. W. A. Cunnington, F.Z.S., 1904—-
1905.—Report on the Copepoda. By Prof. G. O. Sars,
C.M.Z.S.
| Received September 11, 1908. |
(Plates VI.-X XITT.*)
INTRODUCTION.
The Entomostracan fauna of the great inland waters of
Central Africa is still very imperfectly known. Only one of the
big lakes, viz., Victoria Nyanza, has been partly explored in this
respect by German naturalists, the results bemg published in
the great work ‘ Die Thierwelt Deutsch Ost-Atrikas,’ whereas the
other large lakes have remained almost entirely unexplored, so
far as regards the Hntomostraca. During the third Tanganyika
Expedition the conductor, Dr. W. A. Cunnington, made collec-
tions not only in Victoria Nyanza, but also in two other lakes,
viz., Nyasa and Tanganyika. The higher Crustacea contained
in these collections have already been worked out and the results
published in the present journal, for the Brachyura by Dr. Cun-
nington, for the Macrura by Dr. Calman. It has hereby been
proved that Lake Tanganyika differs remarkably in its faunistic
character from the other two lakes, and on the whole exhibits
a much richer and more specialized fauna, as regards these two
groups. Of course, it would be of great interest to know if an
analogous difference also exists as regards the lower Crustacea or
Entomostraca. Material for the solution of this question has
fortunately been procured by Dr. Cunnington, who by the aid
of more or less fine-meshed tow-nets has collected from each
of the three lakes a number of samples chiefly taken at the
* For explanation of the Plates see p. 72.
32 PROF. G. 0. SARS ON THE COPEPODA [Jan. 12,
surface, but in some cases so near the shores as also to contain
some material from the bottom. At the request of Dr. Cunning-
ton, 1 have undertaken the examination of these samples, and
have been much interested in selecting from them the several
forms of Entomostraca therein contained. The Cladocera have
been sent back to Dr. Cunnington, who desired to report on that
group himself, whereas the Copepoda and the Ostracoda will be
worked out by me.
The material which has been placed in my hands for examina-
tion, consists of a large series of tubes containing samples
carefully preserved in formalin, and exactly numbered, both as
to date and locality. To these samples were added some few
smaller tubes containing Copepoda and Ostracoda, selected by
Dr. Cunnington from some of the larger tubes. The greatest
number of samples are derived from Lake Tanganyika. There
are, however, a suflicient number of samples also from the two
other lakes for instituting a comparison of the Entomostracan
fauna of all three lakes.
At the first sight, the samples looked rather unpromising,
some of them being apparently quite devoid of Entomostraca,
and others partly filled up with a compact muddy deposit which
seemed quite unfit for investigation. Yet, by a very careful
microscopical examination of all the tubes, I have succeeded in
bringing to light a considerable number of different forms, both
of Copepoda and Ostracoda, many of them apparently new to
science. Some of the latter are of a quite particular interest,
and will necessitate a renewed discussion about the origin of
the fauna in the lakes. At the close of this Report some general
remarks on this and other questions will be given.
Owing to the great number of new and imperfectly known
forms found in the samples, and which need to be described and
figured, I have found it appropriate to divide my Report into
two parts, the one treating of the Copepoda, the other of the
Ostracoda. To these will be added a smaller paper on some
larval stages of prawns, mostly found in the samples from Lake
Tanganyika. The present part of the Report is wholly devoted
to the Copepoda, which is by far the most richly represented
group of Crustacea in the lakes.
Systematic Nores AND DESCRIPTIONS OF NEW OR IMPERFECTLY
KNOWN SPECIES.
The Copepoda found in the samples belong to the three leading
divisions: Calanoida, Harpacticoida, and Cyclopoida. The first-
named division, as is well known, contains forms which, as a rule,
lead an errant or sub-pelagic life, and of course will chiefly
characterise the surface-plankton of the lakes. On the other
hand, the forms belonging to the second division are exclusively
bottom-dwellers. The third division contains both planktonic
and true bottom-forms.
1909. | OF THE THIRD TANGANYIKA EXPEDITION. 33
I. CALANOIDA.
Of this division five species have been found in the collection,
all of which belong to a single genus
Genus Diapromus Westwood,
Of this genus, as is well known, a very great number of species
have been described, chiefly distributed in the lakes of the
northern hemisphere. South of the equator the genus Loeckella
seems to replace Diaptomus, only a very limited number of species
of the latter genus having hitherto been recorded, viz., 2 from
Victoria Nyanza, 2 from “the Cape Colony, 2 from Australia,
and 6 from South America. Of the five species here recorded,
two have been previously, though rather imperfectly, described,
whereas the other three species are new to science. I give below
short diagnoses of all the five species, and on the accompanying
plates car refully- drawn. habitus- and detail- figures of each.
1, DIAPTOMUS GALEBOIDES, nom. nov. (Plate VI. figs. 1-8.)
Syn.: Diaptomus galebi Mrazek (not Barrois).
Specific Characters.—Frmaur. Body (figs. 1 & 2) moderately
slender, with the anterior division oblong in form, slightly
narrowed both in front and behind, the greatest width occurring
anterior to the middle. Last segment of metasome not defined
from the preceding one, except by a very slight notch on each
side, lateral corners scarcely at all projecting, nearly rectangular,
with a very small and delicate mucroat the tip (see figs. 3 & 4) ;
inner part rounded off, without any mucro. Urosome (fig. 3)
almost quite symmetrical, with the genital segment only very
slightly dilated in front, and carrying on each side an extremely
small and delicate, hair-like point. Last two segments imper-
fectly defined, and combined about half the length of the genital
segment. Caudal rami comparatively short, with the marginal
sete of quite normal appearance. Anterior antenne slender and
elongated, when reflexed exceeding the body by the last 2 or 3
joints (see fig. 2). Last pair of legs (fig. 5) with the inner
ramus quite short, not attaining half the length of the proximal
joint of the outer ; claw of the latter ramus of moderate size and
finely ciliated inside; ; terminal joint only faintly defined at the
base; inner apical seta extending beyond the tip of the claw.
Ovisac (see fig. 1) comparatively small, and containing, as a rule,
only four ova arranged regularly in pairs.
Matz (fig. 6) somewhat smaller than female and of more slender
form. Last segment of metasome with the lateral corners simple,
scarcely produced behind. Urosome narrow cylindric in form,
5-articulate. Right anterior antenna with the projection of the
antepenultimate joint (see fig. 7) very small, not nearly attain-
ing half the length of the penultimate joint, and scarcely at
all curved at the tip; last jomt without any hook at the end.
Last pair of legs (fig. 8) somewhat resembling in structure those
Proc. Zoou. Soc.—1909, No. ITT. 3
34 PROF. G. 0. SARS ON THE COPEPODA [Jan. 12,
in D. galebi, though, on a closer comparison, differmg in some
particulars. Right leg with the 2nd basal joint produced at the
end, on the posterior face, to a short, somewhat faleiform incurved
lamella, inner edge perfectly smooth; proximal joint of outer
ramus terminating outside in an obtuse corner; distal joint
pyriform in shape, with a very delicate scale-like projection on
the posterior face, about in the middle between the apical claw
and the lateral spine, the latter attached much nearer to the base
than to the end of the joint ; apical claw much curved and finely
denticulated inside for some part of its length. Left leg with
the outer ramus distinctly biarticulate and terminating in a well
defined digit accompanied by a small ciliated bristle; imner edge
divided into two rounded and finely ciliated lobules.
Length of adult female 1-30 mm.
Remarks.—Vhis form has been identified by Dr. Mrazek with
the Egyptian species, J). galebi Barrois, apparently owing to a
certain similarity in the structure of the last pair of legs in the
two sexes. I find, however, that the present form differs in so
many other points from the above-named species, as more fully
described by Dr. Richard *, that I have felt justified in regarding
it as specifically distinct, though nearly related to D. galebi. The
specific name here proposed alludes to this near relationship of
the two species.
Occurrence.—This form occurred in great abundance in a
sample taken 25/4/05 off the island Bukoba, western shore of
Victoria Nyanza. The same species has also been recorded pre-
viously by Dr. Mrazek and Dr. von Daday from other localities,
and seems indeed to be one of the most characteristic plankton-
forms of that lake. Together with the usual form, there also
occurred in the same sample a much smaller variety of somewhat
more slender body and with the anterior antenne more elongated,
but otherwise agreeing in all structural details exactly with the
larger form.
2. DIAPTOMUS MIXTUS, sp.n. (Plate VII. figs. 9-17.)
Specific Characters.—FrmMaue. Body (figs. 9 & 10) comparatively
more slender than in the preceding species, with the anterior
division narrow oblong in form and having its greatest width
about in the middle. Last segment of metasome, as in D. gale-
boides, confluent with the preceding one, but having the lateral
corners distinctly projecting, each armed with two well-defined,
sharply-pointed mucros (see fig. 11), the one issuing somewhat
more dorsally. Urosome conspicuously asymmetrical, its terminal
part being more or less turned to left side; genital segment
(fig. 11) gradually widening in front, and armed on each side
with a small, though distinct mucro; last two segments con-
fluent, and combined not attaining half the length of the genital
segment. Caudal sete normal. Anterior antenne very slender
* Revue biologique du nord de la France, tome y. 1892-93.
1909.] OF THE THIRD TANGANYIKA EXPEDITION. 35
and elongated, when reflexed exceeding the body by the last three
joints. Last pair of legs (fig. 12) rather short and stout; inner
ramus about half the length of the proximal joimt of the outer ;
claw of the latter ramus smooth inside ; terminal joint very small,
with the inner apical seta not extended beyond the tip of the
claw.
Mate (fig. 13) with the lateral corners of last segment of
metasome produced backwards and each tipped with a well
defined mucro, the right one more prominent than left. Uvro-
some very slender and more or less turned to right side. Right
anterior antenna (fig. 14) with the middle section rather dilated ;
terminal section, as usual, composed of four joints, the ante-
penultimate one produced at the end anteriorly to arather strong
hook-like projection (fig. 15) considerably exceeding half the
length of the penultimate jot. Last pai: of legs (fig. 16) rather
strongly built ; right leg with the 2nd basal joint produced at the
end, on the posterior face, to a short and broad, securiform plate
projecting into three angular corners (fig. 17); inner ramus
extremely small and rudimentary ; outer ramus with the proximal
joint short and obtusely produced outside ; distal joint very large,
oblong, pyriform in shape, and so attached that it forms nearly a
right angle with the axis of the leg; scale-like projection of
posterior face occurring just inside the lateral spine, the latter
rather slender and attached close to the base; apical claw strong
and much curved. Left leg of a similar structure to that in
D. galeboides.
Length of adult female 1°55 mm.
Remarks.—In the general appearance, the distinctly developed
mucros on the last segment of the metasome, and the con-
spicuously asymmetrical urosome, this form much more resembles
D. galebi than does the preceding species. Yetitis quite certainly
distinct from both these species, as proved by the rather different
structure of the last pair of legs, especially in the male. The
somewhat mixed characters of this species have given rise to the
specific name here proposed.
Occurrence.—A single adult female and some few male speci-
mens of this form were picked up from a sample taken 13/6/04
in Anchorage Bay, southern end of Lake Nyasa. The sample
contained a considerable quantity of a dark muddy deposit, and
thus has apparently been taken from very shallow water. A few
not fully adult. specimens of the same species occurred also in
another sample taken 11/6/04 in that bay.
3. DIAProMUS STUHLMANNI Mrazek. (Plate VIII. figs. 18-24.)
Diaptomus stuhlmanni Mrazek, Die Copepoden Ost-Afrikas,
p. 7, pl. iii. figs. 1-3, 6, 7.
Specific Characters.—Frmaue. Body (figs. 18 & 19) compara-
tively short and stout, with the anterior division somewhat
dilated in front, the greatest width occurring across the cephalic
segment. Last segment of metasome, as in the two preceding
3%
36 PROF. G, 0. SARS ON THE COPEPODA [Jan. 12,
species, confluent with the preceding one; lateral corners rather
projecting, and each terminating im a strong mucro pointing
outwards (fig. 20). Urosome comparatively short and perfectly
symmetrical ; genital segment nearly of equal width throughout
and without any distinct lateral mucros; last two segments
confluent, and combined not nearly attaining half the length of
the genital segment. Caudal rami and sete: normal. Anterior
antenne, when reflexed, scarcely extending beyond the tip of the
caudal rami. Last pair of legs (fig. 21) with the inner ramus
more produced than in the two preceding species, considerably
exceeding half the length of the proximal joint of the outer, and
tipped with two slender sete; claw of outer ramus of moderate:
size and quite smooth ; terminal joint imperfectly defined at the
base, inner apical seta extending to the tip of the claw. Ovisac
comparatively small, with a somewhat varying number of ov:
(from two to eight).
Mate with the antepenultimate joint of right anterior antenna
produced at the end to a slender mucroniform process nearly as.
long as the penultimate joint (fig. 22). Last pair of legs (fig. 23)
with the squamiform expansions of Ist basal joint simple, not.
spiniferous ; 2nd basal joint of right leg without any projection
at the end ; proximal joint of outer ramus terminating outside in
an obtuse corner ; distal joint oblong oval in form, with a small
scale-like projection on the posterior face just inside the lateral
spine, the latter of moderate size and attached about im the
middle of the outer edge; apical claw about twice the length of
the distal joint and somewhat curved in the middle. Left leg
with the outer ramus uniarticulate and somewhat spoon-shaped,
terminating in a small digit accompanied by a short ciliated
bristle ; inner edge evenly curved and finely denticulated; anterior
face with a small ciliated lobule not projecting beyond the inner
edge (fig. 24).
Length of adult female 1-12 mm.
Remarks.—This form has been described, though rather imper-
fectly, under the above name by Dr. Mrazek im the above quoted!
paper. It is nearly allied to the two preceding species, but
evidently specifically distinct, being especially distinguished by
the short and stout form of the body, the projecting lateral
corners of the last segment of the metasome, and the structure of
the last pair of legs in both sexes.
Occurrence.—This form occurred rather plentifully together
with D. galeboides in the above-mentioned sample from Victoria
Nyanza. It was also found in another sample from the same
lake, taken 20/4/05 in about the same region. Dr. Mrazek
records it also from the island Djume.
4, DIAPTOMUS SIMPLEX, sp.n. (Plate VIII. figs. 25-32.)
Specific Characters—FEMALB. Body (figs. 25 & 26) rather short,
with the anterior division regularly oblong-oval in form, gradually
narrowed both in front and behind. Last segment of metasome
1909. | OF THE THIRD TANGANYIKA BXPEDITION. oT
(see fig. 27) wholly coalesced with the preceding one, and having
the lateral parts quite simple, not expanded laterally, each with a
very delicate hair-like point on the outer edge (fig. 28). Urosome
(fig. 27) rather slender and narrow, and perfectly symmetrical ;
genital segment slightly widening in front and without any
lateral mucros; the last two segments confluent, and combined
fully half the length of the genital segment. Caudal rami some-
what more produced than in the three preceding species; marginal
sete normal. Anterior antenne slender and elongated, when
reflexed exceeding the body by the last two joints (see figs, 25 &
26). Last pair of legs (fig. 29) comparatively large; imner ramus
scmewhat exceeding half the length of the proximal joint of the
outer and finely ciliated on the tip; claw of this ramus com-
paratively short and thick; terminal joint extremely minute,
nodiform, with the inner apical seta not extended to the tip of
the claw. Ovisac rather large, broadly rounded in form, and
containing a greater number of ova than ina ny of the preceding
species.
Mate with the antepenultimate joint of right anterior antenna
(fig. 30) produced at the end to a slender spiniform process
slightly curved at the tip, and fully as long as the penultimate
joint. Last pair of legs (fig. 31) with the squamiform expansions
of 1st basal joint, as in JD. stwhlmanni, simple, not spiniferous
2nd basal joint of right leg without any projection at the end
proximal joint of outer ramus terminating outside in an obtuse
corner; distal jot broadly oval in form and without any scale-
like projection of the posterior face; lateral spine comparatively
short and attached much nearer to the end than the base of the
joint ; apical claw of moderate size and only slightly curved. Left
leg with the outer ramus pronouncedly spoon-shaped and uni-
articulate, inner edge evenly curved and minutely spinulose ;
terminal digit very small and partly concealed, accompanying
bristle likewise small; anterior face of the ramus provided with a
small linguiform lamella, finely ciliated at the edge (see fig. 52).
Length of adult female 1:10 mm.
Remarks.— this is a rather small species, and is prominently
distinguished not only from the four preceding species, but also
from most other known Diaptomi, by the simple, not expanded
lateral parts of the last segment of metasome in the female; hence
the specific name here proposed.
Occurrence.—This form occurred rather abundantly in two of
the samples from Tanganyika, the one taken 18/11/04 off Kala,
eastern shore of the lake, the other 1/3/05 off Kaboge, north-
western shore. It was also found occasionally in some other
samples, and seems indeed to be one of the most characteristic
plankton-forms of that lake.
?
?
DIAPTOMUS CUNNINGTONI, sp.n. (Plate TX. figs. 33-42.)
Specific Characters.— Fumaue. Body (figs. 33 & 34) compara-
tively short and stout, with the anterior division somewhat
38 PROF. G. 0. SARS ON THE COPEPODA (Jan. 12,
dilated in its anterior part and but slightly narrowed behind.
Last seement of metasome (see fig. 35) confluent with the pre-
ceding one, and having the lateral parts conspicuously expanded,
each terminating in an acutely produced corner pointing out-
wards, their posterior edge being boldly rounded off and without
any mucro (figs. 36 & 37); left expansion a little larger than
right. Urosome (fig. 35) comparatively short, with the genital
segment slightly asymmetrical and almost of uniform width
throughout; last two segments confluent, and combined not
attaining half the length of the genital segment. Caudal rami
short, with the marginal sete normal. Anterior antennz (see
figs. 33 & 34) remarkably short, when reflexed scarcely extending
beyond the middle of the genital segment. Last pair of legs
(fig. 38) with the inner ramus rather produced, extending almost
to the end of the proximal joint of the outer, and tipped with
two slender sete; claw of the latter ramus comparatively short
and smooth inside; terminal joint very small and imperfectly
defined at the base, its inner apical seta not extending as far as
the claw.
Mats (fig. 39) with the lateral corners of the last segment of meta-
some produced backwards, that on right side the more prominent,
each tipped with a delicate, hair-like mucro. Antepenultimate
joint of right anterior antenna (fig. 40) produced at the end to a
strong spiniform process extending to the end of the succeeding
joint. Last pair of legs (fig. 41) with the squamiform expansions
of lst basal joimt each tipped with a slender hair-like spine;
right leg with the 2nd basal joint simple, without any projection
at the end; proximal joint of outer ramus produced at the outer
corner to an acute spiniform process; distal joint oval in form.
and provided on the posterior face, somewhat in front of the
middle, with a slightly curved spiniform projection ; lateral spine
comparatively short and attached near the end of the joint ;
apical claw rather strong, and abruptly curved in the middle.
Left leg with the outer ramus pronouncedly spoon-shaped and
uniarticulate ; inner edge evenly curved and fringed with short
hairs; apical digit very short and obtuse at the tip; anterior
face of the ramus hollowed and provided with a delicate ciliated
lappet not projecting beyond the inner edge (fig. 42).
Length of adult female 1-25 mm.
Remarks.—Vhis form is at once distinguished from the other
four species here described by the shortness of the anterior
antenne. Moreover, several well-marked differences are found in
the other structural details, as pointed out in the above diagnosis.
I have much pleasure in naming this distinct species in honour
of the distinguished conductor of the Expedition.
Occurrence.—This species is peculiar to Lake Nyasa. I have
found it very abundantly in three of the samples from that lake,
one of them being taken 17/6/04 in Monkey Bay, the other two.
off Karonga, north end of the lake.
1909. ] OF THE THIRD TANGANYIKA EXPEDITION. 39
Il. HARPACTICOIDA.
The forms of this division found in the collection belong to
two distinet genera, both of which have hitherto been regarded
as strictly marine. Hach of these genera belongs to a separate
family, the one to the Diosaccidee, the other to the Cletodide,
both families represented in the sea by numerous genera and
species.
Family DiosaccipD&.
One of the most prominent characters of this family, as indi-
cated by the name, is the presence in the female of two ventral
ovisacs, a feature very seldom met with in the Harpacticoida,
and not found in any of the hitherto described fresh-water forms,
which, like the great majority of marine forms, have only asingle
ovisac. Indeed, this character was formerly regarded as peculiar
to the whole division, in contradistinction to the Cyclopoida,
where constantly two ovisacs are present in the female. The
present family comprises as yet about five distinct genera.
Genus Scuizopera G. O. Sars.
Generic Characters—Body more or less slender, cylindric in
form, with no sharp demarcation between the anterior and
posterior divisions. Cephalic segment of moderate size and
provided in front with a well-developed rostrum, distinctly defined
at the base, and generally of narrow lanceolate form. Epimeral
plates of the three succeeding segments rounded. Last pedigerous
segment without true epimeral plates. Urosome scarcely nar-
rower than the anterior division; genital segment in female
imperfectly divided in the middle. Caudal rami more or less
produced, generally somewhat lamellar, each with a strong spine
outside accompanied by a slender seta, and with another delicate
seta arising from the upper face ; tip truncated and carrying three
sete, the innermost extremely small, the outermost much shorter
than the middle one, which generally is very strongly developed.
Anterior antennze in female comparatively short, 8-articulate ;
Ath joint somewhat produced at the end and carrying a slender
band-like sensory appendage, terminal part consisting of four
joints, the last much the longest; those in male hinged in the
usual manner. Posterior antenne comparatively short and stout,
geniculate; outer ramus small, biarticulate. Mandibles with the
palp well developed, biramous. Maxilla and maxillipeds of usual
structure. First pair of legs more or less distinctly prehensile ;
outer ramus 3-articulate, and generally shorter than the inner ;
the latter somewhat varying in structure in the different species,
consisting of three or only two joints, the first of which is the
largest and provided inside at some distance from the end with
a strong ciliated seta. Natatory legs, as a rule, very slender,
with both rami 3-articulate and provided inside with a very
limited number of spiniform sete; inner ramus of 2nd pair of
40 PROF. G. 0. SARS ON THE COPEPODA [Jan. 12,
legs in male transformed, carrying near the tip outside a strong
spiniform appendage. Last pair of legs comparatively small,
distal joint with generally six very unequal and partly spiniform
marginal sete, inner expansion of proximal joint rather short and
provided with four spiniform setze; these legs in male, as usual,
still smaller and with the number of sets reduced. Ovisacs in
female narrow and closely juxtaposed, each containing a very limited
number of ova. In some cases arranged in a single row.
Remarks.—This genus was established by the present author
in the year 1905 *, to comprise a species, S.. longicauda G. O. Sars,
found in a brackish lagoon on the Chatham Islands. It was,
however, subsequently + withdrawn, as I found it somewhat
ditticult to distinguish it exactly from the much diversified genus
Amphiascus G.O. Sars. Having, however, now become acquainted
with a number of new species agr eeing with that originally
observed in the more essential char aoa Se I am disposed to restore
the present genus, which seems to form a quite natural group.
None of the Norwegian species of Amphiascus can properly be
referred to it. On the other hand, a well-marked species oceurs
in the Caspian Sea, and another species I have recently found in
a sample from the brackish lake, Birket el Qurun, in Egypt.
In the present collection I have determined no less than eight
different species, to be described below.
6. SCHIZOPERA INOPINATA, sp.n. (Plate X. figs. 43-58.)
Specific Characters.—FrMaun. Body (figs. 43 & 44) moderately
slender and of the usual subcylindric form, with the anterior
division longer than the posterior. Rostrum (see fig. 45) rather
prominent, narrow lanceolate in form, and somewhat curved at the
tip. Last caudal segment (see fig. 55) shorter than the preceding
one, and having ‘ne. anal opercle - very small and perfectly smooth.
Caudal rami about the length of that segment and rather diver-
gent, being gradually narrowed distally, inner edge straight and
very finely ciliated; spine of outer edge attached about in the
middle, dorsal seta somewhat nearer the tip, the latter narrowly
truncated and armed above the insertion of the apical setee with
a transverse row of four small denticles. Anterior antenne (see
fig. 45) comparatively slender, though not nearly attaining the
length of the cephalic segment. Posterior antenne (fig. 46)
remarkably short and stout, with the outer ramus (fig. 47) very
narrow. Oral parts (figs. 48 & 49) of the structure characteristic
of the genus. First pair of legs (fig. 50) with the outer ramus
about the length of the Ist joint of the inner, and having the
last joint considerably longer than the other two 05 inner ramus
distinctly 3-articulate, w ith the last two joints incurved and
combined scarcely attaining half the length of the 1st; last joint
armed at the tip with a Bentler claw-like spine and two unequal
* Zool. Jahrbiicher, Bd. xxi. Heft 4, p. 383.
+ Account of the Crustacea of Norway, vol. v. p. 148.
1909. ] OF THE THIRD TANGANYIKA EXPEDITION. 4]
sete. Natatory legs (figs. 51-53) with no seta inside the Ist
joint of the rami. ‘Last pair of legs (fig. 54) with the distal joint
oval quadrangular in form, proximal seta of outer edge rather
strong and spiniform, innermost seta but one likewise strongly
developed; inner expansion of proximal joint somewhat narrowed
and extending to about the middle of the distal joint. Ovisacs
narrow oblong, with the ova partly arranged in a single row.
MAtLe with the anterior antenne (fig. 56) distinctly hinged.
Inner ramus of 2nd pair of legs (fig. 57) transformed in the usual
manner. Last pair of legs (fig. 58) very small, with only four
sete on the distal joint and two sete on the end of the inner
expansion of proximal joint.
Length of adult female 0-45 mm.
Remarks.—This was the first species which came to my sight,
and I have therefore given to it the above specific name. From
the typical species, S. longicauda, it is easily distinguished by the
comparatively shorter urosome and the somewhat differ ent shape
of the caudal rami. Moreover, there are some differences to be
found in the structure of the legs. The present species is also
rather inferior in size.
Occurrence.—This form was found occasionally in two of the
samples from Tanganyika, the one taken 28/9/04 at Mbete, south
end of the lake, the other, 13/10/04, at Sumbu, south-western
border.
7. SCHIZOPERA VALIDIOR, sp. n. (Plate XI. figs. 59-63.)
Specific Characters.—Frmaue. Body (fig. 59) on the whole
more strongly built than in the preceding species, with the
integuments of coarser consistency. Rostrum somewhat less
produced, but of a similar narrow lanceolate form. Caudal rami
(fig. 63) rather large and broad, pronouncedly lamellar, and of
oval form, less dive ergent than in SS. inopinata; inner edge
distinctly curved and densely hairy, spine of outer edge somewhat
shorter than in that species, and attached much nearer to the
end than to the base of the ramus; dorsal seta, on the other hand,
issuing rather in front of the middle; end of the ramus without
any denticles above. Anterior antenne (fig. 61) of a similar
structure to that in S. inopinata, though perhaps a little shorter.
Posterior antenne less robust. First pair of legs (fig. 61) with the
outer ramus about as in the last-mentioned species, Inner ramus
distinctly 3-articulate; Ist jomt not fully as long as the outer
ramus, the last two joints comparatively more elongated than in
S. inopinata, and combined considerably exceeding half the length
of the lst; last joint, as in that species, armed on the tip with a
slender claw-like spine and two unequal sete. Natatory legs
with a well-developed seta inside the Ist joint of the inner ramus
in the two posterior pairs. Last pair of legs (fig. 62) with the
distal joint of a more regular oval form than in S. inopinata
and somewhat conically produced at the tip; proximal seta of
outer edge very long and abruptly deflexed ; innermost seta but
42 - PROF. G, O. SARS ON THE COPEPODA [Jan. 12,
one likewise more elongated than in that species ; inner expansion
of proximal joint comparatively larger, extending somewhat
beyond the middle of the distal jomt. Ovisaes (see fig. 59) broader
than in that species, and containing a greater number of ova
arranged in a double row.
Length of adult female 0°49 mm.
Remarks.—This form is closely allied to S. inopinata, but of
somewhat larger size and more robust form of body. It also
differs somewhat in the shape of the caudal rami and in the
structure of the legs, as indicated in the above diagnosis.
Occurrence.—Only a single female specimen of this form has
come to my notice. It was found in one of the two samples
in which the preceding species occurred, viz., that from Sumbu.
8. SCHIZOPERA CONSIMILIS, sp. n. (Plate XI. figs. 64-67.)
Specific Characters.—VEMALE. Body (fig. 64) considerably more
slender than in the two preceding species, and somewhat atten-
uated behind. Rostrum about as in S. validior. Urosome
almost attaining the length of the anterior division, last segment
somewhat shorter and narrower than the preceding one and
having the anal aperture smooth. Caudal rami (fig. 67) con-
siderably produced, exceeding in length the anal segment, and
rather divergent, being nearly of uniform width throughout,
inner edge straight and finely ciliated, spine of outer edge
attached much nearer to the end than to the base of the ramus ;
dorsal seta likewise issuing somewhat behind the middle; tip
transversely truncated and armed above the insertion of the
apical setee with a transverse row of about seven small denticles.
Antenne about as in S. validior. First pair of legs (fig. 65)
comparatively shorter and stouter; outer ramus with the last
joint shorter than the other two; inner ramus distinctly 3-arti-
culate, with the Ist joint a little longer than the outer ramus,
last two joints comparatively short, and combined scarcely
attaining one-third of the length of the Ist; last joint, as in the
two preceding species, armed with a slender claw-like spime and
two unequal sete. Natatory legs of a similar structure to those
in S. validior. Last pair of legs (fig. 66) with the distal joint
comparatively small, rounded oval in form; marginal sete about
as in S. validior ; inner expansion of proximal joint rather short,
not extending to the middle of the distal joint. Ovisacs oblong
in form, with the ova arranged in a double row.
Length of adult female 0-51 mm.
Remarks.—This is perhaps the form which comes nearest to
the typical species, S. longicauda. On a closer comparison it is,
however, found to differ in some particulars, for instance in the
shape of the caudal rami, and also somewhat in the structure of
the legs, proving it to be in reality specifically distinct.
Occurrence.—Some few specimens of this form were found,
partly in the same sample in which S. validior occurred, partly
in another sample from Tanganyika, taken 29/10/04 in Vua
1909. ] OF THE THIRD TANGANYIKA EXPEDITION. 43
Harbour, western shore of the lake. Finally a single female
specimen was found in the above-mentioned sample from Victoria
Nyanza,
9. SCHIZOPERA UNGULATA, sp.n. (Plate XI. figs. 68-71.)
Specific Characters.—Frmaue. Body (fig. 68) rather slender,
sublinear in form or very slightly attenuated behind. Rostrum
long and prominent. Last caudal segment a little shorter than
the. preceding one, but scarcely narrower ; anal opercle very small
and perfectly smooth. Caudal rami (fig. 71) rather darge and
broad, conspicuously lamellar and somewhat divergent; inner
edge cre! and densely hairy; spine of outer edge ainachied
nearer to the end than to the base of the ramus; “danse seta.
originating about in the middle; end narrowly truncated and
without any denticles above. Anterior antenne rather slender,
but of usual structure. First pair of legs (fig. 69) with the outer
ramus comparatively short and stout, having the last joint
longer than the others; inner ramus Sirol built, but only
composed of two joints, the Ist considerably longer: than the
outer ramus and slightly curved, the 2nd quite short and abruptly
incurved, carrying on the tip a strong claw and a comparatively
short seta, accompanied by a small hair-like bristle. Natatory
legs rather strongly built, but scarcely differing in their structure
from those in the two preceding species. Last pair of legs (fig. 70)
with all the marginal sete, except that issuing from the tip of
the distal joint, remarkably strong and spiniform, being also
much shorter than in the three preceding species ; inner expansion
of proximal joint extending almost to the tip of the distal
joint.
Length of adult female 0°50 mm.
Remarks.—This species is chiefly distinguished from the pre-
ceding ones by the rather different structure of the Ist pair of
legs, the inner ramus of which is composed of only two joints
and is more prominently prehensile than in those species. It
also differs somewhat in the shape of the caudal rami and the
structure of the last pair of legs.
Occurrence.—Only a single specimen of this form, a female
with well developed ovaria, but without ovisacs, has come to my
notice. It was found in the same sample in which S. validior
occurred.
10. ScHIZOPERA MINUTICORNIS, sp.n. (Plate XII. figs. 72-77.)
Specific Characters.—FrmMaun. Body (fig. 72) very slender and
narrow, sublinear in form, with the posterior division fully as
long as the anterior. Rostrum (see fig. 73) acutely produced.
Last caudal segment a little shorter and narrower than the pre-
ceding one; anal opercle very small and quite smooth. Caudal
rami (fig. 77) pronouncedly lamellar and of oval form, being only
slightly divergent; inner edge evenly curved and very finely
ciliated, spine of outer edge not very strong and attached at a
d4 PROF. G. O. SARS ON THE COPEFODA [Jan, 12,
short distance fiom the end of the ramus; dorsal seta issuing
about in the middle, apical sete comparatively short. Anterior
antenne (see fig. 73) unusually small, though composed of the
normal number of joints. First pair of legs (fig. 74) with the
outer ramus of normal structure; mner ramus almost twice as
long, but composed of only two joints of about equal length,
the distal one linear in form and scarcely at all incurved, carrying
on the tip a slender claw-like spine and a single still more slender
seta. Natatory legs (fig. 75) with the seta at the inner corner of
the terminal joimt of outer ramus very small and rudimentary ;
[st joint of inner ramus in 2nd pair of legs without any seta
inside. Last pair of legs (fig. 76) with the distal joint com-
paratively short, rounded oval in form, marginal sete of the
usual appearance ; inner expansion of proximal joint extending
about as far as the distal joint.
Length of adult female 0°46 mm.
Remarks.—This species is at once distinguished by the
unusually small anterior antenne, a character which has given
vise to the specific name here proposed. It also can be distin-
euished by the anomalous structure of the Ist pair of legs.
Occurrence.—Of this form also only a solitary specimen has
come to my notice, a fully grown female with well-developed
ovaria, but without ovisacs. It was found in the same sample as
the preceding species.
11. SCHIZOPERA SPINULOSA, Sp. n. (Plate XII. figs. 78-81.)
Specific Characters.—Frmaur. Body (fig. 78) not particularly
slender, with the posterior division much shorter than the
anterior and slightly tapering behind. Rostrum of the usual
shape. Last caudal segment somewhat shorter than the preceding
one, and exhibiting above, on each side of the anal area, an
oblique row of about five small spinules; anal opercle perfectly
smooth. Caudal rami (fig. 81) broadly oval in form and _ pro-
nouncedly lamellar ; inner edge evenly curved and armed with a
row of slender spinules continued across the end of the ramus
above the insertion of the apical sete; spine of outer edge rather
strong and attached at a short distance from the end; dorsai seta
issuing at about the middle. Anterior antennz comparatively
slender and of quite normal structure. First pair of legs (fig. 79)
somewhat resembling in structure those in S. wnagulata, the inner
ramus being composed of only two very unequal joints; the Ist
long and slender, considerably exceeding in length the outer
ramus; the 2nd quite short and armed on the tip with a strong
claw and a comparatively short seta accompanied by a small hair.
Natatory legs about as in the four preceding species. Last pair
ot legs (fig. 80) resembling in shape those in S. minuticornis,
though wanting one of the small marginal spines on the distal
joint.
Length of adult female 0°43 mm.
1909. ] OF THE THIRD TANGANYIKA EXPEDITION, 4D.
Remarks.—The peculiar armature of the anal segment and of
the caudal rami will suffice for distinguishing at once this species
from any of the preceding, and it 1s from this character that
the specific name here proposed is derived. In the structure of
the Ist pair of legs it differs conspicuously from the type.
Occurrence. —A single female specimen of this form was
found in the above-mentioned sample taken in Tanganyika at
Mbete.
SCHIZOPERA FIMBRIATA, sp. n. (Plate XII. figs. 82-86.)
Specific Characters.—FrMAue. Body (fig. 82) moderately slender
and slightly attenuated behind. Rostrum well defined and of
the usual narrow lanceolate form. Last caudal segment fully as
long as the preceding one, and, as in S. spinalosa, armed above
on each side of the anal area with an oblique row of small
spinules; anal opercle well developed, semilunar, and fringed
with a regular row of about 16 similar spinules. Caudal rami
(fig. 86) considerably shorter than the anal segment and of oval
form; inner edge somewhat curved at the base; spine of outer
edge short and thick, curved outwards, and attached somewhat
nearer to the end than to the base; dorsal seta likewise issuing
behind the middle; upper face of each ramus armed with three
transverse rows of extremely fine spinules, the Ist crossing the
base, the 2nd extending from the inner edge to the dorsal’ seta,
the 3rd occurring just above the insertion of the apical setee.
Anterior antennee comparatively short, but composed of the
normal number of joits. First pair of legs (fig. 83) with both
rami comparatively short, last joint of outer ramus not attaining
the length of the middle one; inner ramus composed of only two
joints, the Ist about the length of the outer ramus, the 2nd
scarcely more than one-third as long and armed on the tip with
a claw-like spime and a slender seta accompanied by a small
hair-like bristle. Natatory legs (fig. 84) with unusually short
rami, but otherwise of normal structure. Last pair of legs
(fig. 85) likewise shorter than usual; distal joint scarcely longer
than it is broad, and, as in S. spinulosa, only provided with five
marginal sete; inner expansion of proximal joint extending
about as far as the distal joint.
Length of the specimen examined 0-44 mm.
Remarks.—The most characteristic feature of this species is
undoubtedly the structure of the anal opercle, which, as in the
species of the genus WVitocra, is fringed with a regular row of
spinules, and it is indeed from this character that the specific
name here proposed is derived. Moreover, the peculiar shape of
the outer spine of the caudal rami is rather characteristic and
affords another easily observable specific mark,
Occurrence.—A single, apparently not yet fully grown female
specimen of this form was found in a sample from Lake Nyasa,
taken 13/6/04 in Anchorage Bay, south end of the lake.
46 PROF. G. 0. SARS ON THE COPEPODA [Jan. 12,
13. ScHIZOPERA SCALARIS, sp. n. (Plate XIII. figs. 87-92.)
Specific Characters.—Maue. Body (fig. 87) slender and elon-
gated, with rather coarse integuments and all the segments very
sharply marked off from each other, giving it a “ sealariform
appearance. Rostrum well defined at the base, but of a some-
what unusual form, being very broad, lamellar, ain terminating
in an obtuse point. Segments of the Acrior division each with
a small nodiform prominence laterally. Last caudal segment
shorter and narrower than the preceding one and without any
spinules dorsally; anal opercle perfectly smooth. Caudal rami
(fig. 92) much produced and very narrow, sublinear in form;
inner edge straight and quite smooth; spine of outer edge
moderately strong and attached at a short distance from the end ;
dorsal seta likewise issuing far behind the middle; median apical
seta very strong and elongated. Anterior antenne (fig. 88) of
moderate size, and hinged in the usual manner. First pair of
legs (fig. 89) ria a highly chitinized, obtusely pointed projection
issuing from the inner corner of the 2nd basal joint, in addition
to the usual spine; outer ramus of quite normal structure; inner
ramus distinctly prehensile, 3-articulate, Ist joint long and
slender, considerably exceeding in length the outer ramus, the
other two joints quite short and somewhat imperfectly separated ;
last joint armed at the tip with a strong curved claw anda slender
seta accompanied by a small hair-like bristle. Natatory legs
well developed and of the structure characteristic of the genus ;
inner ramus of .2nd pair of legs (fig. 90) transformed in whe usual
manner. Last pair of legs (fig. 91) with the distal joint com-
paratively small and sores fusiform in shape, with only five
very unequal marginal sete ; inner expansion of proximal joint
short and ‘amounted at the tip, which carries two thickish juxta-
posed sete ciliated in their outer part.
Length of the specu examined 0-49 mm.
Remaris. arance this form looks very
different from the giahen nse, and by its pronouncedly sealari-
form body, the broad rostral plate, and the slender and narrow
caudal rami, 1 much more resembles some species of the genera
Laophonte and Cletodes, The anatomical examination has,
however, proved it to be a genuine member of the present
genus.
Occurrence.—The above-described male specimen, the only one
that has come to my notice, was found in a sample from Tangan-
yika, taken 24/2/05 at Baraka, north-western shore of the lake.
Family CLETODID &.
This family is chiefly distinguished from the Zaophontide, to
which it bears some resemblance, by the non-prehensile Ist pair
of legs. The type of the family i is the genus Cletodes Brady, to
which im recent times several other genera have been added, all
from the sea.
1909.] OF THE THIRD TANGANYIKA EXPEDITION. 47
Genus Inyorninus Lilljeborg.
Generic Characters. e or less cylindric in form, with
rather thin and flexible integuments and the segments sharply
marked off from each other. Cephalic segment large, and pro-
jecting in front to a rounded rostral plate not defined at the
base. Urosome attenuated behind, with the genital segment
distinctly divided in the middle. Caudal rami more or less
produced, not lamellar, with one of the apical sete very strong,
spiniform. Anterior antennze in female composed of only five
joints, thickly clothed with partly spiniform sete; those in male
distinctly hinged. Posterior antenne rather strong, with the
distal jomt armed with coarse claw-like spines; outer ramus
small, uniarticulate. Mandibular palp likewise uniarticulate.
Maxille and maxillipeds comparatively short and stout. First
pair of legs only slightly differing from the succeeding ones, and
not prehensile ; inner ramus in this and the two succeeding pairs
biarticulate and much smaller than the outer, that of the 4th
pair quite rudimentary, uniarticulate; 5rd pair of legs in male
slightly transformed. Last pair of legs comparatively small, with
the distal joint well defined or confluent with the proximal one;
inner expansion of the latter broad, but not projecting. A single
ovisac present in female.
Remarks.—This genus was established in the year 1902 by
Prof. Lilljeborg, to comprise a small Harpacticoid, /. flexibils
Lilljeborg, found by him off the shores of the Baltic in the neigh-
bourhood of Stockholm. I have myself met with the same species
near Christiania, in shallow bays of the Fjord, and thus have
had an opportunity of examining it more closely. Prof. Lilljeborg
refers this genus to the subfamily Nannopodine of Brady, which,
according to that author, comprises the two genera Vannopus
and Platychelipus. The first of these genera is still very. imper-
fectly known, but is apparently referable to the family Cletodide.
The second genus, in my opinion, is very different, and ought to be
referred to the family Laophontide. In any case, the present genus
seems to me to be a genuine Cletodid, exhibiting, as it does, all
the essential characters of that family.
14. ILYOPHILUS PERPLEXUS, sp.n, (Plate XIII. figs. 93-104.)
Specific Characters.— FrmMae. Body (figs. 93 & 94) rather short
and stout, more or less curved, and gradually attenuated from
before backwards, with no sharp demarcation between the
anterior and posterior divisions. All the segments of the body
sharply defined, and having the hind edge somewhat raised but
quite smooth. Cephalic seement very large and broad, with a
slight dorsal depression about in the middle, and with the lateral
parts boldly curved anteriorly. Rostral projection evenly rounded
at the tip and densely clothed with fine hairs (see fig. 95). The
four succeeding segments comparatively short and obtusely pro-
duced later: ally. Urosome almost as long as the anterior division ;
48 PROF, G. 0. SARS ON THE COPEPODA - [Jan. 12,
genital segment very distinctly divided in the middle ; last segment
fully as long as the preceding one and slightly widening at the
end; anal opercle small, but somewhat prominent, and perfectly
smooth. Caudal rami about twice as long as they are broad, sub-
linear in form and slightly diverging, each carrying on the tip a
strong spiniform seta tipped with a slender bristle and accom-
panied on each side by a very small hair-hke bristle; seta of outer
edge likewise small, hair-hke, and attached about in the middle ;
dorsal seta issuing somewhat nearer to the end. Anterior
antenne (fig. 95) rather strong and curved, with the Ist joint
very thick, though scarcely longer than the 2nd; 38rd joint
carrying at the end anteriorly the usual sensory filament, which
is rather small; the last two joints, forming together the terminal
part, very unequal, the Ist quite short, the 2nd somewhat lamellar
and armed with two strong spines and several thin flexible sete.
Posterior antenne (fig. 96) with five strong claw-like spines on the
distal joint, increasing in length distally ; outer ramus very small
and only provided with three sete at the tip. Mandibular palp
(fig. 97) likewise smaller than in the typical species. Maxille
(fig. 98) and anterior maxillipeds (fig. 99) of about the same
appearance as in that species. Posterior maxillipeds (fig, 100),
however, comparatively less powerful, Ist pair of legs (fig, 101)
with the inner corner of 2nd basal joint conically produced and
tipped with a strong spine; terminal joint of outer ramus armed
with three slender spines and a single seta issuing from the inner
corner; inner ramus scarcely longer than the Ist joint of the
outer. The two succeeding pairs of legs (fig. 102) only differing
from the 1st in the want of a spine at the inner corner of the
2nd basal joint, and in the terminal joint of outer ramus being
somewhat broader and provided with one or two additional sete.
4th pair of legs (fig. 103) with one of these sete shortened and
spiniform ; inner ramus quite rudimentary, consisting of a single
small, nodiform joint tipped with a slender seta. Last pair of
legs (fig. 104) rather small, with the distal and proximal jomts
wholly confluent, forming together a short and broad lamella
produced outside to a conical process, to which a slender hair-lke
bristle is attached; posterior edge of the lamella irregularly
indented and fringed with seven thickish, finely ciliated sete.
Length of adult female 0°49 mm.
Remarks.—Vhe above-described form unquestionably is referable
to the genus Jlyophilus, as defined by Prof. Lilljeborg, but is
quite certainly specifically distinct from the typical species
I. flewibilis, differing, as it does, in some pomts very markedly.
Thus the caudal rami are much more produced, and the last pair
of legs are distinguished by the complete coalescence of the
two joints of which these legs are originally composed. It is also
rather inferior in size.
Occurrence —Only a single female specimen of this remarkable
form has come to my notice. It was picked up from some muddy
deposit contained ina sample from Tanganyika, taken 13/10/04
1909. ] OF THE THIRD TANGANYIKA EXPEDITION. 49
at Sumbu, western shore of the lake. From the same sample
also several of the above described species of the genus Schizopera
were derived.
Tit. CYCLOPOIDA.
The Cyclopoida found in the collection are referable to two
particular sections of that extensive division, viz., the Gnatho-
stomata and the Peecilostomata. The first of these sections com-
prises the typical free-living Cyclopoida, in which the oral parts
are normally developed. To the second section, on the other
hand, are referred forms which lead a more or less parasitic life,
and in which therefore the oral parts have been transformed in
accordance therewith, though not to such an extent as in a third
section, not represented in the collection, the Siphonostomata.
Sect. Gnathostomata.
Family CycLopipa.
Genus Cyctors Miiller.
By far the great majority of Copepod-species found in the
collection belong to this extensive and widely distributed genus.
No less than 20 different species have been determined, 16 of
which have proved to be new to science.
The distinction of the species of this genus is in some cases
determined with no small difficulty. For, whereas some of them
are at once recognised, there are other species which are so
closely related, that a very careful examination is needed for
distinguishing the one from the other. For the discrimination
of nearly allied species, much stress has generally been laid on
the structure of the rudimentary last pair of legs. The exami-
nation of these diminutive appendages is, however, by no means
easy, and cannot be effected without a very careful dissection of the
specimens. There are several other characters, which are much
more easily accessible for examination and consequently of more
practical value. Among them may be mentioned the mutual
relation in length of the caudal sete. As this relation has proved
to be perfectly constant in all individuals of the same species, and
admits of being examined at once, without any dissection of the
specimens, it affords one of the practically best characters for the
distinction of nearly-related species. Especially is the mutual
relation between the innermost and outermost apical seta in
many cases very characteristic,
Owing to the great number of species comprised within this
genus, it will probably in future be found advisable to effect a
subdivision of it into several nearly allied genera. It is well
known indeed that Dr. Schmeil, in his excellent account of the
fresh-water Cyclopide, has arranged the European species in
several groups, and in recent times some attempts have been made
Proc. Zoo. Soc.—1909, No. LV. 4
dU PROF. G, 0. SARS ON THE COPEPODA (Jan. 12,
to raise these groups to distinct genera or subgenera. In the
present paper I find it, however, appropriate to retain the old
generic name Cyclops for all the species here recorded, though
they are enumerated in a definite succession according to their
affinity.
15. Cyctors teucKartr Claus. (Plate XIV. figs. 105-107.)
Oceurrence.—This truly cosmopolitan species was found in
several of the samples from all three lakes. In some of the
samples from Tanganyika it occurred in great abundance, and
may, together with Diaptomus simplex, form a chief part of the
surface-plankton of that lake. In a sample from Nyasa, taken
13/6/04 in the Anchorage Bay, a smaller variety occurred not:
unfrequently. For comparison with the two succeeding species,
I give on the accompanying plate a figure of a typical female
specimen from Victoria Nyanza, together with two detail-
figures.
16. Cycrors emtnt Mrazek. (Plate XIV. figs. 108-112.)
Cyclops emini Mrazek, Die Copepoden Ost-Afrikas, p. 4, pl. u.
figs. 1-3, 5, 6, 8.
Specific Characters.— FEMALE. Body (fig. 108) rather slender,
with the anterior division oblong-oval in form and shghtly widening
anteriorly, front narrowly truncated. Last pedigerous segment
small, not produced laterally. Urosome slender and narrow,
considerably exceeding half the length of the anterior division ;.
genital segment only very slightly widening in front. Caudal
rami (fig. 112) much longer than the anal segment and rather
narrow, being somewhat divergent ; seta of outer edge attached a.
little behind the middle ; innermost apical seta more than twice as.
long as the outermost, but considerably shorter than the outer
mediate one; dorsal seta unusually long and slender, extending
beyond the innermost apical seta. Anterior antenne very long
and slender, almost attaining the length of the whole anterior
division, and 17-articulate. Posterior antenne (fig. 109) likewise
anusually slender, with the terminal joint narrow linear in form,
and the penultimate joint only provided with four anteriorly-
curving sete. Natatory legs with the rami comparatively long
and slender ; inner ramus of fourth pair (fig. 110) with the two
apical spines very unequal, the inner one rather slender, the outer
scarcely more than one-third as long, both distinctly denticulate.
Last pair of legs (fig. 111) very small, distal jomt narrow oblong
in form, and having the lateral spine longer than the apical seta.
Ovisacs comparatively small and somewhat divergent, each with
a rather limited number of ova.
Leneth of adult female 0°79 mm.
Remarks.—This form has been described, though somewhat
imperfectly, under the above name by Dr. Mrazek. It is closely
allied to the European species, C. oithonoides G.O. Sars, exhibiting
a very similar external appearance, though at once distinguished
1909. | OF THE THIRD TANGANYIKA EXPEDITION. 51
by the comparatively shorter innermost caudal seta. In C. oithon-
oides this seta is about the same length as the outer mediate one,
whereas in the present species it is scarcely more than two-thirds
as long. The extraordinary length of the dorsal seta has also
been noted by Dr. Mrazek. He has, however, not been aware of
the rather striking difference from most other species in the
structure of the posterior antennz. According to its organisation,
this species, ike C. lewckarti, may be regarded as a true limnetic
or planktonic form.
Occurrence.—I have only found this form in one of the samples,
viz., that from Bukoba, Victoria Nyanza. It did not occur in
any ‘abundance, Dr. Mrizek has recor ded it also from the island
Djuma in the same lake.
17. CYCLOPS NEGLECTUS, nom. nov. (Plate XIV. figs. 113-
Ute)
Syn.: Cyclops hyalinus Richard (not Rehberg).
- oithonoides Mrazek (not G. O. Sars).
Specific Characters.—Frmaun. Body (fig. 113) much shorter
and stouter than in the preceding species, with the anterior
division regularly oval in form and the front evenly rounded.
Last pedigerous segment small, not produced later ally. Urosome
about half the length of the anterior division, and having the
genital segment slightly widening in front. Caudal rami (fig. 117)
very short, searcely exceeding in length the anal segment, and
almost twice as long as they are broad ; seta of outer edge attached
at about the posterior third of the ramus; apical setae compara-
tively short, immermost one not attaining half the length of the
outer mediate seta and scarcely twice as long as the outermost ;
dorsal seta of moderate length. Anterior antenne not nearly
attaining the length of the anterior division of the body, and
17-articulate. Posterior antennze (fig. 114) of normal appearance,
with about nine anteriorly-curving setze on the penultimate joint
Natatory legs with the rami less Slender than in C. emini; inner
ramus of fourth pair (fig. 115) with the apical spines very unequal
in size. Last pair of legs (fig. 116) resembling those in the said -
species, though having the lateral spine of the distal joint com-
paratively shor ter. Ovisacs not very lar ge, oval in form, and only
containing a limited number of ova.
Length of adult female 0°73 mm.
Remarks.—This form is closely allied to the European species
CO. hyalinws Rehberg, and indeed Dr. Richard, and formerly
also I myself, identified it with that species. It is, however, on
a closer comparison, at once distinguished by the much shorter
innermost caudal seta. Whereas in Oh yalinus this seta is about
the same length as the outer mediate one, it is in the present form
searcely more than half as long. Dr. Mrazek has identified this
form with C. oithonoides G. 0. Sars, to which species it in reality
bears only a very remote resemblance.
Ae
t
r
e
Y
2 PROF. G. 0. SARS ON THE COPEPODA [Jan. 12,
Occurrence.—Of this species specimens were found in samples
from all three lakes; but only in one of them, taken 13/6/04 in
the Anchorage Bay, Nyasa, it occurred in any abundance.
Distribution. —Senegal at Rufinesque (Richard), Sumatra
(G. O. Sars); ? Germany (Lichard).
18. CYCLoPs TENELLUS, sp.n. (Plate XIV. figs. 118-127.)
Specific Characters.—FEMALE. Body (fig. 118) rather slender,
with very thin and fragile integuments. Anterior division of
body regularly oval in form, with the cephalic segment very large
and somewhat truncated in front. Last pedigerous segment very
small. Urosome slender and narrow, exceeding half the length
of the anterior division ; genital segment only very slightly dilated
in front. Caudal rami (fig. 127) about twice as long as they are
broad and scarcely divergent; seta of outer edge attached nearly
in the middle, innermost apical seta very small, scarcely longer
than the outermost, inner mediate seta very slender and almost
twice as long as the outer, dorsal seta likewise rather much pro-
duced. Anterior antenne (fig. 119) slender and elongated,
attaining almost the length of the anterior division of the body,
though composed of only twelve articulations clothed with slender
bristles, some of them of quite an unusual length. Posterior
antenne (fig. 120) likewise rather slender, with a very limited
number of sete (5) on the penultimate joint. Both pairs of maxilli-
peds (fig. 121) unusually produced. Natatory legs well developed ;
terminal joint of outer ramus in the two anterior pairs (figs. 122 &
123) provided with five slender spines and five sete, in 3rd pair
(fig. 124) with only four spines; Ist joimt of same ramus, except
in the Ist pair, without any seta inside; 4th pair (fig. 125) with
both rami very slender and devoid of the seta inside the Ist joint ;
apical spines of inner ramus very unequal in size. Last pair of
legs (fig. 126) extremely small, though apparently of normal
structure, apical seta the longest.
Length of adult female 0°38 mm.
Remarks.—This is perhaps the smallest of all the species of
Cyclops, and is also distinguished rather conspicuously from the
other known forms both in its external appearance and the
structure of the appendages. It therefore appears somewhat
difficult to decide to which group of the genus Cyclops it should
more properly be referred. I place it provisionally here, next to
the three preceding species, to which it exhibits a certain similarity
as to the general habitus. To judge from its very thin and
pelucid integument and the delicate structure of the appendages,
it seems to be a true limnetic form.
Occurrence.—I have only found this tiny species in two of the
samples from Tanganyika, one taken 24/2/05 at Baraka, north-
western shore of the lake, the other 1/3/05 at Kaboge, in the
same region of the lake. Most of the specimens were more or
less damaged, owing to the extremely fragile structure of their
bodies.
1909. | OF THE THIRD TANGANYIKA EXPEDITION. 53
19. CycLors aLErpus (Jurine). (Plate XV. figs. 128-130.)
Occurrence.—-Some few specimens of this well-known species,
most of them of the male sex, were found in a sample from Lake
Nyasa, taken 11/6/04 in the Anchorage Bay. I give on the
accompanying plate a figure of a male specimen together with two
detail-figures.
Disiribution.— Kurope, Asia, North and South America,
Australia, Hawaii Islands.
20. CYCLOPS ATTENUATUS, sp. n. (Plate XVI. figs. 131-138.)
Specific Characters—Frmaue. Body (fig. 131) comparatively
slender and attenuated behind, with the anterior division oval in
form and evenly rounded in front. Last pedigerous segment
slightly produced on each side. Urosome somewhat exceeding
half the length of the anterior division, and gradually tapering
behind; genital segment only shghtly dilated in front. Caudal
rami (fig. 138) rather produced, exceeding in length the last two
segments combined, and of narrow linear form, being not at all
divergent; seta of outer edge somewhat remote from the end;
innermost apical seta very thin and about twice the length of the
outermost, which is somewhat spiniform; the two middle sete
long and slender, the mimner one exceeding the outer by rather
more than one-third of its length and almost attaining half the
length of the whole body. Anterior antennee (fig. 152) compara-
tively short, not nearly attaining the length of the cephalic
segment, and composed of only ten articulations. Posterior -
antennee (fig. 133) of normal structure, with six anteriorly-curving
sete on the penultimate joint. Natatory legs (figs. 134-136)
short and stout, with both rami bi-articulate, the last two joints
being wholly coalesced. Last pair of legs (fig. 137) with the
proximal joint not defined, its seta issuing from the side of the
segment itself ; distal joint very small, narrow conical in form and
tipped with a single seta. Ovisacs small, rounded, each as a rule
containing only two or three globular ova.
Length of adult female 0:65 mm.
Remarks.—This form is nearly allied to the Kuropean species,
C. varicans G. O. Sars, having, like the latter, both rami of the
natatory legs bi-articulate, and the last pair of legs partly confluent
with the pertaining segment. It differs, however, in the some-
what more slender form of the body, the more produced caudal
rami, and in the anterior antennz being composed of only 10
articulations, whereas in C. varicans they are distinctly 12-
articulate.
Occurrence.—Several specimens of this form were picked up
from the muddy deposit contained in one of the samples from
Tanganyika, taken 13/10/04 at Sumbu, south-western shore of
the lake, and two specimens, male and female, were found in
another sample, taken 28/9/04 at Mbete, south end of the lake.
D4 PROF. G. 0. SARS ON THE COPEPODA (Jan. 12,
21. Cycrops varicans G. O. Sars. (Plate XVI. figs. 139, 140.)
Occurrence—A single female specimen of this form was
found in a sample from Lake Nyasa, taken 13/6/04 in the
Anchorage Bay, and another somewhat smaller specimen in a
sample from Tanganyika, taken 28/9/04 at Mbete, south end of
the lake.
Distribution.—Kurope, New Zealand (4. O. Sars).
22. CYCLOPS EXIGUUS, sp.n. (Plate XVI. figs. 141, 142.)
Specific Characters.—FEMALE. Body (fig. 141) rather narrow,
with the anterior division oblong-oval in form and obtusely pro-
duced in front. Last pedigerous segment with the lateral parts
slightly prominent and each tipped with a slender bristle.
Urosome slender, considerably exceeding half the length of the
anterior division; genital segment only very slightly dilated in
front. Caudal rami (fig. 142) not attaiming the length of the
last two segments combined, of nearly uniform width throughout,
and scarcely at all divergent; seta of outer edge attached some-
what behind the middle, innermost apical seta very thin and fully .
twice as long as the outermost, the two middle setz of moderate
length and having the cilia of their proximal part much coarser
than those of the distal part, the inner one exceeding the outer
by about one-third of its length. Anterior antenne rather short,
and composed of only 11 articulations. Legs of a similar
structure to those of the two preceding species.
Length of adult female 0°53 mm.
Remarks.—Vhe present form is closely allied to the European
species, C. bicolor G. O. Sars, but is of smaller size and more
narrow form of body. It also differs in the shorter anterior
antennz and in the less produced caudal rami.
Occurrence.—Only a single female specimen of this form has
come to my notice. It was found in a sample from Tanganyika,
taken 28/9/04 at Mbete, south end of the lake.
23. CYCLOPS CUNNINGTONI, sp.n. (Plate XVII. figs. 143-150.)
Specific Characters—Fermaue. Body (fig. 143) comparatively
short and robust, with the anterior division rather broad and
pronouncedly depressed, having its greatest width somewhat
behind the middle. Cephalic segment very large, and gradually
narrowed in front to an obtusely conical point. Lateral parts of
the three succeeding segments somewhat expanded, but with the
hind corners rounded off. Last pedigerous segment forming on
each side a short obtuse prominence tipped with a slender ciliated
seta. Urosome rather thick, about half the length of the anterior
division and slightly tapering behind; genital segment somewhat
dilated in its anterior part. Caudal rami (fig. 150) not nearly
attaining the length of the last two segments combined and
slightly widening at the end; seta of outer edge attached some-
wh at behind the middle; innermost apical seta only slightly longer
1909. ] OF THE THIRD TANGANYIKA EXPEDITION. DD
than the outermost, which is spiniform; the two middle sete
rather coarse, almost spiniform, and of moderate length; the inner
one exceeding the outer by about one-third of its length, dorsal
seta not much produced. Anterior antenne (fig. 144) compara-
tively short and thick at the base, consisting of 12 articulations
densely clothed with slender curved sete especially on the proximal
part. Posterior antenne (fig. 145) with the seta attached to the hind
corner of the basal joint rather short; penultimate joint provided with
nine anteriorly-curving sete, and about the length of the terminal
joint. Natatory legs (figs. 146-148) short and stout and, as in
the three preceding species, having both rami bi-articulate. Last
pair of legs (fig. 149) quite rudimentary, each consisting of only
a very small conical joint tipped with a single seta, the basal
joint being wholly confluent with the pertaining segment. Ovi-
sacs of moderate size, oblong oval in form, and somewhat divergent,
each containing from ten to sixteen ova.
Length of adult female 0°86 mm.
Remarks.—This is a v ery distinct and easily recognisable form,
being especially distinguished from the three preceding species,
with which it a grees in the structure of the legs, by the short and
robust body, the broad, pronouncedly depressed anterior division,
and the conically produced cephalic segment. It is indeed one
of the most characteristic endemic forms of Lake Tanganyika,
and I have therefore thought it right to associate with it the
name of the distinguished conductor of the Expedition.
Occurrence.—1 have found this pretty form in four of the
samples from Tanganyika. In one of these samples, the same
which contained the above-described species, C. atéenwatus, it
occurred rather frequently, more rarely in the three other samples,
‘Of these one was taken 12/12/04 at Karema, eastern shore of the
lake, another 1/3/05 at Kaboge, north-western shore, and a third
28/9/04 at Mbete, south end of the lake.
24. CYCLOPS PACHYCOMUS, sp. n. (Plate XVII. figs. 151-156.)
Specific Characters.—FEMALE. Body (fig. 151) considerably
more slender than in C. cwnningtoni, with the anterior division
less expanded and the cephalic segment obtusely rounded in front.
Last pedigerous segment about as in that species. Urosome some-
what exceeding half the length of the anterior division and
eradually tapering behind. Caudal rami (fig. 156) somewhat
narrower than in C. cunningtoni, though scarcely longer, and not
at all diverging ; seta of outer edge attached about in the middle,
innermost apical seta scarcely longer than the outermost, the two
middle sete rather slender and elongated, the inner one almost
twice as long as the outer and exceeding half the length of the
whole body. Anterior antenne (fig. 152) very short and robust,
and composed of only 11 articulations thickly covered with
very coarse, almost spiniform set, several of which are distinctly
ciliated. Posterior antenne (fig. 153) likewise shorter and stouter
than in the preceding species. Natatory legs of a very similar
56 PROF. G. 0. SARS ON THE COPEPODA [Jan. 12
structure to that in C. cunningtoni, except that the spines on both
rami of the 4th pair (fig. 154) are shorter and blunt at the tip.
Last pair of legs (fig. 155) exhibiting the same rudimentary
structure as in that species.
Length of adult female 0°62 mm.
Remarks.—Though closely allied to the preceding species, this
form is at once distinguished by the less expanded anterior
division of the body, the different shape of the cephalic segment,
the more slender caudal sete, and finally by the shorter and
thicker anterior antennee, which moreover are only 11-articulate,
and have the sete of the anterior edge and upper face remarkably
coarse. The specific name here proposed is derived from this
last character.
Occurrence.—Some few female specimens of this form were
found in two of the samples from Tanganyika, the one taken
28/9/04 at Mbete, south end of the lake, the other 13/10/04 at
Sumbu, south- weston shore.
25, CYCLOPS SEMISERRATUS, sp. n. (Plate XVIII. figs. 157-
169.)
Specific Characters. — Frmaur. Body (fig. 157) moderately
slender, with the anterior division regularly oval in form, the
greatest width occurring about in the middle. Cephalic segment
lar ge and obtusely truncated in front. Last pedigerous segment
with the lateral parts shghtly expanded and clothed w ith fine
hairs. Urosome somewhat exceeding half the length of the
anterior division; genital segment “comparatively caltont and
distinctly dilated ‘at the base, carrying on each side a slender
bristle (see fig. 167). Caudal rami (fig. 169) long and slender,
considerably exceeding in length the last three segments com-
bined, and of narrow linear form, slightly widening at the tip;
outer edge armed for about half its length with a row of very
small denticles, seta of this edge small and attached close to the
end somewhat dorsally, innermost apical seta only shightly longer
and much thinner than the outermost ; middle sete rather slender
with the cilia of uniform appearance throughout, the inner one
exceeding the outer by rather more than one-thir d of its length
and abruptly bent outwards at the middle, Anterior antenne
(fig. 158) comparatively slender, considerably exceeding the
cephalic segment in length, and composed of 12 articulations.
Posterior antenne (fig. 159) of moderate length, and having the
seta of the basal joint well developed. Oral parts (figs. 160-163)
and natatory legs (figs. 164-166) built on the same type as in
C. serrulatus Fischer. Last pair of legs (figs. 167, 168) as in that
Species, each composed of a single small, somewhat lamelliform
joint, trilobate at the end and carrying two slender bristles and
at the inner corner a ciliated spine, the latter, however, much
smaller than in C. serrulatus. Ovisacs regulaxly oval in form and
only very slightly divergent.
Length of adult female 0:86 mm.
1909. | OF THE THIRD TANGANYIKA EXPEDITION. 57
Remarks.—This form, like the six succeeding species, belongs to
the group of Cyclops for which C. serrulatus Fischer is the type,
and which has proved to contain several well-defined species
formerly generally regarded as only varieties of the said species.
The present form is nearly allied to C. macruroides Lilljeborg,
differing, however, in some particulars, for instance in the arma-
ture of the caudal rami and the mutual relation in length of the
apical seta, so it may be more properly regarded as specifically
distinct.
Occurrence.—This form occurred very abundantly in-one of the
samples from Tanganyika, taken 19/9/04 at Niamkolo Island,
south end of the lake. It was not found in any of the other
samples.
26. CYCLOPS LEVIMARGO, sp.n. (Plate XIX. figs. 170-175.)
Specific Characters.—Frmaue. Body (fig. 170) comparatively
more robust than in the preceding species, with the anterior division
shghtly dilated in front. Last pedigerous segment and urosome
about as in that species. Caudal rami (fig. 175) of a similar
narrow linear form, but with the outer edge perfectly smooth,
without any trace of the regular row of denticles found in that
species ; innermost apical seta scar cely longer than the outermost,
the two middle setz comparatively shorter than in Q. semiser-
ratus and quite uniformly ciliated, the inner one only exceeding
the outer by one-fourth of its length and slightly bent outwards
at the middle. Anterior antenne (fig, 171) much shorter than
in that species, scarcely attaining the length of the cephalic
segment, though composed of the same number of articulations.
Posterior antennee (fig. 172) likewise shorter and stouter. Fourth
pair of legs (fig. 173) with the spines on the outer ramus com-
paratively ‘short and blunt at the tip. Last pair of legs (fig. 174)
with the spine of the inner corner small, though perhaps a little
longer than in the preceding species. Ovisacs comparatively small
and scarcely divergent.
Length of adult female 0°85 mm.
Remarks.—The present form differs conspicuously from the
preceding one by the much shorter anterior antenne and by
the absolute absence on the outer edge of the caudal rami of the
regular series of denticles generally found in the species belonging
to this group. It is from this last character that the specific
name here proposed is derived.
Occurrence.—Vhis form also was only found in one of the
samples from Tanganyika, but in this rather abundantly. The
sample was taken 12/12/05 at Karema, eastern shore of the lake.
27. CycLops aANcusTUS, sp. n. (Plate XIX. figs. 176-180.)
Specific Characters.—FrmMaue. Body (fig. 176) very slender and
narrow, with the anterior division oblong in form and obtusely
truncated in front. Last pedigerous segment with the lateral
parts only slightly expanded. Urosome attaining about two-
58 PROF, G. 0, SARS ON THE COPEPODA [Jan. 12,
thirds of the length of the anterior division; genital segment
conspicuously dilated at the base. Caudal rami (fig. 180) ex-
ceedingly slender and elongated, almost attaining the length of
the remaining part of the urosome ; outer edge perfectly smooth,
seta of this edge attached at only a short distance from the end
and accompanied at the base in front by a transverse row of four
small denticles ; innermost apical seta about the length of the
outermost ; the two middle sete rather slender and uniformly
ciated, the inner one exceeding the outer by about one-third of
its length. Anterior antenne 12-articulate and of moderate
length, being about as long as the cephalic segment, their outer
joints (fig. 177) conspicuously longer than in ©. levimargo.
Natatory legs of normal structure. Last pair of legs (fig. 179)
extremely small, with the spine of the inner corner very short.
Ovisacs about as in the two preceding species.
Length of adult female 0°82 mm.
Remarks.—This form at the first sight looks very like the
Huropean species, C. macrurus G. O. Sars. On a closer com-
parison, however, it is found to differ in the comparatively more
slender anterior antenne, and more particularly in the structure
of the caudal rami. In the European species the seta of the
outer edge is much more remote from the end, and in front of it
four denticles occur, which do not form a transverse row, as in
the present species, but are arranged along the edge. Moreover,
the innermost apical seta is considerably longer than the outer-
most, and the spine of the inner corner in the last pair of legs
more fully developed.
Occurrence.—Some few specimens of this form were found in a
sample from Tanganyika, taken 18/11/04 at Kala, eastern shore
of the lake. One female and two male specimens of apparently
the same species occurred in a sample from Lake Nyasa, taken
13/6/04 in the Anchorage Bay.
28. CYCLOPS RARISPINUS, sp. n. (Plate XIX. figs. 181, 182.)
Specific Characters. —FumMaue. Body (fig. 181) comparatively
short and stout, with the anterior division ovoid in form and
narrowly truncated in front. Last pedigerous segment with the
lateral parts somewhat produced and clothed with fine hairs.
Urosome slightly exceeding half the length of the anterior
division, and having the genital segment rather much dilated at
the base. Caudal rami (fig. 182) much shorter than in the three
preceding species and also less narrow, scarcely widening at the
end: outer edge armed with a short row of about eight small
denticles not extending to the middle of the ramus, inner edge
finely ciliated in its proximal half ; innermost apical seta nearly
twice as long as the outermost; the two middle set rather slender
and uniformly ciliated, the inner one exceeding the outer by rather
more than one-third of its length. Anterior antenne moderately
slender and distinctly 12-articulate. Legs apparently of normal
structure. Ovisacs comparatively small and scarcely divergent.
1909. | OF THE THIRD TANGANYIKA EXPEDITION. 59
Length of adult female 0°80 mm.
Remarks.—In the form and armature of the caudal rami this
form somewhat resembles a species recorded by Dr. Mrazek from
small ponds at Bukoba, Victoria Nyanza, and named C. stuhl-
manni. To judge from the figure given by that author, this
form is, however, much more slender in shape, and the anterior
antenne are composed of only 10 articulations, whereas in the
present form they are distinctly 12-articulate, as in all the other
species belonging to this group.
Occurrence.—Only a single female specimen of this form has
come to my notice. It was found in a sample from Tanganyika,
taken 18/11/04 at Kala, eastern shore of the lake.
29, CYCLOPS AGILOIDES, sp.n. (Plate XX. figs. 183-188.)
Specific Characters.—FrMAuE. Body (fig. 183) rather slender,
with the anterior division gradually widening anteriorly, front
evenly rounded. Last pedigerous segment short, but rather
broad. Urosome slender, considerably exceeding half the length
of the anterior division ; genital segment somewhat dilated at the
base. Caudal rami (fig. 188) of moderate length, about as long
as the last two segments combined, and slightly divergent; outer
edge armed with a row of very small denticles, which in the
proximal part of the ramus are somewhat withdrawn from the
edge and become so minute that they easily may escape attention ;
innermost apical seta nearly twice as long as the outermost ; the
two middle sete of moderate length and uniformly ciliated, the
inner one exceeding the outer by rather more than one-third
of its length. Anterior antenne rather slender, considerably
exceeding in length the cephalic segment, and 12-articulate, the
last three joints (fig. 184) with a distinct smooth longitudinal
keel. Posterior antenne (fig. 185) of quite normal structure.
Fourth pair of legs (fig. 186) with the spines of both rami finely
denticulate, the apical ones of the inner ramus slightly unequal
in size. Last pair of legs (fig. 187) with the spine of the inner
corner well developed and coarsely denticulate.
Length of adult female 0°90 mm.
Remarks.—This form is closely allied to the European species,
C. agilis Koch (= C. varius Lilljeborg), but differs in some
particulars, especially as regards the form and armature of the
caudal rami, so as more properly to be regarded as specifically
distinct.
Occurrence.—A. single female specimen (that described above)
of this form was found in the above-mentioned sample from
Victoria Nyanza, and two other specimens of apparently the same
species weve derived from Tanganyika, the one from a sample
taken 19/9/04 at Niamkolo, south end of the lake, the other in a
sample taken 13/10/04 at Sumbu, south-western shore.
30. CycLops EUACANTHUS, sp.n. (Plate XX. figs. 189-192.)
Specific Characters—FrmMae. Body (fig. 189) comparatively
60 PROF. G. 0. SARS ON THE COPEPODA (Jan. 12,
slender, with the anterior division oblong-oval in form and nar-
rowly truncated in front. Last pedigerous segment with the
lateral parts slightly produced and finely hairy. Urosome ex-
ceeding half the length of the anterior division, and having the
genital segment somewhat dilated in front. Caudal rami (fig. 192)
about half the length of the remaining part of urosome and
somewhat divergent, outer edge armed with a comb-like row of
rather strong denticles extending almost to the base of the ramus ;
innermost apical seta very thin and quite naked, being more than
twice as long as the outermost, the latter pronouncedly spiniform
and pointing almost straight outwards, its anterior edge fringed
with fine spinules, posterior edge smooth ; middle sete slender and
uniformly ciliated, the inner one exceeding the outer by rather
more than one-third of its length. Anterior antenne scarcely as
long as the cephalic segment, and 12-articulate; last and penulti-
mate joints (see fig. 190) each provided with a finely denticulated
longitudinal keel. Fourth pair of legs (fig. 191) with the spines
attached to the terminal joint of both rami unusually large,
lanceolate in form, with a dense fringe of delicate spmules on
each side, Last pair of legs not examined. Ovisacs of moderate
size and oval in form, slightly divergent.
Length of adult female 0°79 mm.
Remarks.-—This form is especially distinguished by the very
conspicuous comb-like series of denticles on the cuter edge of the
caudal rami, the strong spiniform outermost apical seta, and
finally by the unusual development of the spines attached to the
terminal joint of both rami in the fourth pair of legs. It is from
this latter character that the specific name here proposed is
derived.
Occurrence-—Two female specimens of this form had been
mounted by Dr. Cunnington, together with some other Ento-
mostraca, on a slide, which was kindly sent to me for examination.
Accor ding to the label, they were collected from the Lofu river,
Tanganyika.
31. CYcCLoPs CILIATUS, sp.n. (Plate XX. figs. 193, 194.)
Specific Characters.— FEMALE. Body (fig. 193) resembling, as to
the general form, that of the preceding species, though perhaps
somewhat less slender. Caudal rami (fig. 194) comparatively
narrower, with the denticles of the outer edge smaller, inner edge
clothed throughout its whole length with delicate, somewhat
distant cilia; innermost apical seta distinetly ciliated and only
slightly exceeding in length the outermost; the latter moderately
strong, with the outer edge minutely denticulated, the imner
clothed with long cilia; middle sete rather slender, with the cilia
of the proximal part coarser and more distant than those of the
distal part, the inner one almost twice as long as the outer.
Anterior antennz more slender and elongated than in the pre-
ceding species, with no keel on the outer joints. Fourth pair of
legs with the spines of quite normal appearance. Last pair of
legs (not figured) with the spine of the inner corner very small.
1909. | OF THE THIRD TANGANYIKA EXPEDITION. 61
Length of adult female 0°86 mm.
Remarks.—TVhe present form is distinguished from the pre-
ceding one, to which it bears some resemblance in its external
appearance, by the comparatively more elongated anterior
antenne, the quite normal structure of the spines attached to
the rami of the fourth pair of legs, and finally by the somewhat
different form and armature of the caudal rami. The distinctly
ciliated inner edge of these rami is another character by which
the present species is distinguished, and which has given rise
to the specific name here proposed.
Occurrence.—Two female specimens of this form (of which one
was dissected) were found in a sample from Tanganyika, taken
28/9/04 at Mbete, south end of the lake.
32. CYCLOPS OLIGARTHRUS, sp.n. (Plate X XI. figs. 195-202.)
Specific Characters.— FEMALE. Body (fig. 195) rather strongly
built and somewhat depressed, with the anterior division ovoid in
form, frontal part conically produced. Last pedigerous segment
short but rather broad, and having the lateral parts densely
clothed with stiff hairs (see fig. 201). Urosome somewhat robust,
with the genital segment considerably dilated at the base. Caudal
rami (fig. 202) rather produced, about equalling in length the
last three segments combined, and slightly diverging; dorsal face
armed, in front of the very small seta of the outer edge, with an
obliquely transverse row of very small denticles; all the apical
setae pronouncedly spiniform and without true cilia, the innermost
and outermost ones of about equal size; inner mediate seta nearly
three times as long as the outer, both clothed in their distal part
with very minute denticles; dorsal seta quite short. Anterior
antenne (fig. 196) unusually short and compact, scarcely ex-
ceeding half the length of the cephalic segment, and composed
of only six articulations densely clothed with coarse sets, some
of which exhibit a comb-like series of thin spinules on the one
edge, one of these sets issuing from the end of the first joint
being particularly strong and extending along the outer part of
the antenna to its very tip. Posterior antenne (fig. 197) likewise
unusually short and stout, with the outer two joints imperfectly
separated and the seta of the basal joint very long. Natatory
legs (figs. 198-200) resembling in structure those in C. fimbriatus
Fischer, the middle joint of the inner ramus being very large and
acutely produced at the outer corner, with the edge in front of
the latter coarsely spmulose. Last pair of legs (fig. 201) each
consisting of a single somewhat irregular joint armed with three
strong spines, the outermost of which is much the largest and
distinctly denticulate, pointing outwards, so as to project on each
side of the pertaining segment (see fig. 195). Ovisacs compara-
tively small, and each containing only a very limited number
of ova.
Length of adult female 0°69 mm.
Remarks.—The nearest ally of this form is unquestionably the
European species, C. jimbriatus Fischer, with which it agrees in
62 PROF. G. O. SARS ON THE COPEPODA [Jan, 12,
the more general characters. Yet it is quite certainly specifically
distinct from that species, exhibiting as it does a number of
well-marked differences, of which may be named the spimiform
character of the caudal sete, the coarse spines with which the last:
pair of legs are armed, and the unusually small number of joints
in the anterior antenne. No other true Cyclops has such a small
number of joints in these antenne, and it is only in the genus
Halicyclops that a similar composition of the anterior antenne is
met with. The specific name here proposed refers to this last
character.
Occurrence.—Some few specimens of this remarkable form were
found in a sample from Tanganyika, taken 28/9/04 at Mbete,
south end of the lake. 'Two specimens (male and female) occur red
in another sample, taken 12/12/04 at Karema, eastern shore of
the lake.
33. CYCLOPS COMPACTUS, sp. n. (Plate X XI. figs. 203, 204.)
Specific Characters.—FEMALE. Body (fig. 203) of a very compact
structure, with the anterior division broadly oval in form and
pronouncedly depressed. Cephalic segment gradnally contracted
anteriorly, with the frontal part narrowly rounded. Lateral parts
of the three succeeding segments somewhat expanded. Last
pedigerous segment short, “but rather broad. Urosome con-
siderably exceeding half the length of the anterior division and
only very slightly “tapering behind; last segment comparatively
short, and clothed laterally with fine spinules. Caudal rami
somewhat produced, equalling in length the last two segments
combined ; inner edge straight and perfectly smooth, outer edge
armed somewhat behind the middle with two or three small
denticles, seta of this edge very small and attached near the end
somewhat dorsally; tip obliquely truncated, with the innermost
seta much smaller than the outermost ; middle setze well-developed
and clothed for some part of their length with small spinules,
the inner one much elongated, considerably exceeding half the
length of the body and fully tivice as long as the outer. Anterior
antenne comparatively short and not much dilated at the base,
being composed of 10 articulations. Last pair of legs (see
fig. 304) each composed of a short joint extending laterally and
carrying three unusually large recurved sete, the longest of which
seme. as far as the second ‘eninge segment, all the sete clothed
with scattered spinules.
Length of adult female 0°75 mm.
Remarks.—This form 1s closely allied to the European species,
C. phaleratus Koch, but is easily distinguished by the much more
produced caudal rami and by the great development of the setz
attached to the last pair of legs.
Oceurrence.—A single female specimen of this form oceurred
in the same mounted slide which contained the two specimens of
the above-described species, C. ewacanthus,
1909. ] OF THE THIRD TANGANYIKA EXPEDITION. 63.
34, CYCLOPS DUBIUS, sp.n. (Plate XXI. figs. 205-207.)
Specific Characters.—FrmMaue. Body (fig. 205) rather slender,
with the anterior division oblong-oval in form and obtusely
rounded in front. Last pedigerous segment (see fig. 206) with
the lateral parts rounded off and clothed with unusually long and
delicate hairs. Urosome considerably exceeding half the length
of the anterior division and somewhat tapering behind; genital
segment only slightly widened in front. Caudal rami (fig. 207)
long and slender, sublinear m form and slightly divergent, con-
siderably exceeding in length the last two segments combined,
outer edge perfectly smooth with a small seta near the end;
innermost apical seta small, being much shorter than the outer-
most, which is spiniform; middle sete of moderate length and
uniformly ciliated, the inner one about twice as long as the outer ;
dorsal seta not much produced. Anterior antenne rather short,
not nearly attaining the length of the cephalic segment and
composed of 12 articulations. Last pair of legs (see fig. 206)
each consisting of a single somewhat lamellar joint extended
laterally, and provided with three comparatively short spiniform
sete.
Length of adult female 0°62 mm.
Remarks.—The exact relation of this form to the other known
species is as yet somewhat doubtful, as the solitary specimen
examined has not been dissected. Yet, in spite of the perfectly
smooth outer edge of the caudal rami, the short anterior antenne,
and the laterally extended last pair of legs, it may probably belong
to the group for which C. serrulatus is the type. In any case
it may be regarded as a rather abnormal form.
Occurrence.—The above-described specimen was found in a
sample from Nyasa, taken 15/6/04 in the Anchorage Bay, south
end of the lake.
Sect. Peecilostomata.
Fam. ERGASILIDG4.
This family comprises a peculiar group of peecilostomatous
Cyclopoida, which in some respects forms as it were a transition
to the true parasites, Caligoida and Lerneoida. The adult
females, which in some cases appear rather much deformed, are
found firmly attached. by the aid of their clawed posterior
antenne, to the gills of several species of freshwater fishes. The
copulation of the sexes in all probability takes place before
the fixation of the female, in the last free stage of the latter.
We know of two distinct genera of this family, viz. Argasilus
Nordman and 7hersites Pagenstecher. To these a third genus is
here added.
Genus ERGASILOIDES, n. g.
Generic Characters.—Body of female (in last free stage) cyclo-
poid in shape, subdepressed in front, and attenuated behind; in
64 PROF. G, 0. SARS ON THE COPEPODA [Jan. 12,
male of more slender form. Head distinctly defined from meta-
some and very large, more or less scutiform and without any
rostrum, Segments of metasome rapidly diminishing in size, the
fifth or last very small, almost obsolete. Uvrosome short, and
consisting in female of only two, in male of three segments ;
genital segment in both sexes large and tumid, that of male
having the postero-lateral corners produced and tipped with a
small spine. Caudal rami short, each carrying on the inner
corner a strong seta, which in some cases is bifid, outer corner
provided with three much smaller sete, one of which is extended
outwards. Anterior antenne of a similar structure in the two
sexes, comparatively short and stout, 5- or 6-articulate, and
densely clothed with delicate sete. Posterior antenne transformed
into strong prehensile organs, larger in female than in male.
Oral area produced in the form of a short tube issuing from the
ventral face of the head behind, and provided inside with two
pairs of incurved appendages (mandibles and maxille), outside
with a small lamella (maxilliped), which in male is transformed
into a well-developed prehensile organ terminating in a clawed
hand. Natatory legs with the rami more or less meurved and
clothed at the end with slender ciliated seta : those of the fourth
pair much smaller than the others, with the number of joints in
the outer ramus reduced. Last pair of legs extremely small and
rudimentary, knob-like, with a single small seta on the tip.
Adult stage of female still unknown.
Remarks.—Vhis new genus is chiefly distinguished from H7rga-
silus, to which it bears a close resemblance, by the reduced
number of segments in the urosome of both sexes. In the corre-
sponding stage of Hrgasilus the urosome is composed in the
female of four well-defined segments and in the male of five such
segments. Moreover, the very rudimentary condition of the last
pair of legs and of the segment to which they ave attached, is
yather characteristic. Three different species of this genus have
been determined.
35, ERGASILOIDES MEGACHEIR, sp.n. (Plate XXII. figs. 208—
Specitic Characters.—FEMALE in last free stage. Body (figs. 208 &
209) comparatively short and stout, pronouncedly depressed, and,
viewed dorsally, subpyriform in outline. Head very large and
expanded, almost quadrangular in form ; dorsal face with a well-
marked transverse fold at about the posterior third of its length,
and moreover exhibiting two small but sharply defined areas, the
one of circular form and occurring in front of the middle, the
other cordiform and placed near the posterior edge; frontal edge
of the head transversely truncated, postero-lateral corners only
slightly prominent and rounded. The four anterior segments of
metasome with the lateral parts slightly produced backwards but
obtusely rounded at the end. Last segment almost wholly
concealed. Urosome scarcely exceeding in length one-third of
1909. | OF THE THIRD TANGANYIKA EXPEDITION. 65
the metasome; genital segment much dilated, rounded oval in
form. Caudal rami about the length of the last segment, with
the seta of the inner corner simple and pointing straight behind.
Anterior antennee (fig. 210) consisting of six well-defined articu-
lations gradually diminishing in size. Posterior antenne (fig. 211)
very largely developed ; propodus twice as long as the basal joint
and oblong in form, being scarcely narrowed distally, anterior
edge with a thin hyaline border not fully extending to the base ;
dactylus about half the length of the propodus and somewhat
twisted, its terminal claw comparatively short, with a recurved
denticle inside. Oral parts (see figs. 212-215) exhibiting the
structure characteristic of the genus. Natatory legs (figs. 216—
218) with both rami distinctly 3-articulate, except the outer
ramus of 4th pair (fig. 218), which is only biarticulate. Last
pair of legs (fig. 219) extremely small.
MALE (fig. 220) smaller than female and of considerably more
slender form. Head much less expanded and rounded in front,
its dorsal face without any distinct sculpturing. Segments of
metasome evenly rounded laterally. Urosome more slender than
in female and about half the length of the metasome. Anterior
antenne only 5-articulate. Posterior antenne (fig. 221) much
smaller than in female and of simpler structure. Maxillipeds
(fig. 222) well developed, with the dactylus very slender.
Length of female in last free stage 0°62 mm., of male
0°55 mm.
Remarks.—This form is especially distinguished by the great
size and peculiar structure of the prehensile posterior antenne in
the female, which character has given rise to the specific name
here proposed.
Occurrence.—Three female specimens (in the last free stage)
and some few immature ones were found in a sample from
Tanganyika, taken 13/10/04 at Sumbu, south-western shore of
the lake.
36. ERGASILOIDES MACRODACTYLUS, sp. n. (Plate XXIII.
figs. 223, 224.)
Specific Characters. —F EMAL (in last free stage). Body (fig. 223)
resembling in its general shape that of the preceding species,
though perhaps a little more slender. Head very large and
expanded, subquadrate in outline, with the dorsal face sculptured
in a similar manner to that in the preceding species, frontal
margin transversely truncated, postero-lateral corners distinctly
projecting, subangular. Urosome with the genital segment less
tumid, inner caudal seta simple. Anterior antenne composed
of only five articulations. Posterior antenne (fig. 224) very
long and slender ; propodus fully twice as long as the basal joint
and attenuated distally, with no hyaline border; dactylus re-
markably elongated, almost attaining the length of the propodus,
with the terminal claw slender and perfectly smooth. Legs of
apparently the same structure as in the preceding species.
Proc. Zoou. Soc.—1909, No. V. 5
66 PROF. G. 0, SARS ON THE COPEPODA [Jan. 12,
Length of female (in last free stage) 0°50 mm.
Remarks.—Vhis form is nearly allied to the preceding one, but
of smaller size, and moreover easily distinguished by the rather
different shape of the prehensile posterior antenne, the dactylus
of which is unusually long and slender. The specific name here
proposed refers to this last character.
Occurrence.—A solitary female specimen of this form was
found in the same sample in which the preceding species
occurred.
37. ERGASILOIDES BREVIMANUS, sp. n. (Plate XXIII. figs. 225—
231.)
Specific Characters.— FEMALE (in last free stage). Body
(fig. 225) somewhat fusiform in shape, the head bulging con-
siderably in its posterior part and gradually tapering anteriorly,
front obtusely rounded. Dorsal face quite smooth without any
trace of the peculiar sculpture found in the two preceding species.
Segments of metasome evenly rounded laterally. Urosome
(fig. 230) of a similar structure to that in the two preceding
species. Caudal rami, however, distinguished by the seta of the
inner corner being bifid, or divided near the base into two some-
what unequal prongs, the outer one being the longer. Anterior
anton composed of only five articulations, Posterior antennee
(fig. 226) much shorter and stouter than in the two preceding
species, with the propodus scarcely longer than the basal joint
and quite simple; dactylus strong and somewhat dilated at the
base, its terminal claw evenly curved and perfectly smooth.
Natatory legs (figs. 227-229) with the outer two joints of the rami
confluent.
MALE (fig. 231) resembling in its external appearance that
of H. megacheir, but at once recognisable by the bifid inner
caudal sete.
Length of female 0°62 mm., of male 0-47 mm.
Remarks.—This form may be easily distinguished from the
two preceding species by the different shape of the head in the
female, the peculiar bifid inner caudal setz, and more particularly
by the much shorter and stouter posterior antenne in the female,
which latter character has given rise to the specific name here
proposed.
Occurrence.—Two or three females and one male specimen of
this form were found ina sample from Tanganyika, taken 28/9/04
at Mbete, south end of the lake. <A single female specimen
occurred in a sample from Nyasa, taken 13/6/04 in Anchorage
Bay.
Genus Ercastnus Nordman.
38. ERGASILUS sp.
Occurrence.—A single female specimen (in the last free stage)
of a genuine Hr gusilus, with distinctly 4-articulate urosome and
1909. | OF THE THIRD TANGANYIKA EXPEDITION. 67
the last pair of legs developed in exactly the same manner as in
the Kuropean species, is present in a mounted slide containing
some other Entomostraca, which, according to the label, are
from Lake Nyasa. The specimen has not yet been examined
in detail, and is therefore only mentioned here. Dr. Mrazek
also records a species of this genus, probably the same as that
here mentioned, from Victoria Nyanza.
GENERAL REMARKS.
The working out of the Copepoda of the Tanganyika Ex-
pedition has involved no. small diticulty and trouble, both as
regards the selection of the specimens from the samples, and the
examination and determination of the species, some of which,
especially of the genus Cyclops, are so closely related to each
other and to European species, that a very minute and careful
examination has been needed to make out their true relationship.
Yet, I think that the labour thereon bestowed may not have
been in vain. For the final results of my examination have
turned out to be on the whole very satisfactory, and have indeed
far surpassed the expectation at first entertained. I hope there-
fore that the present Report will furnish a not unimportant
contribution both to the exact definition of species, and to the
general characterisation of the fauna in the three great Central
African lakes.
The number of Copepod-species examined and mentioned in
the present Report amounts to no less than 38 in all, belonging to
six different genera. Of these species the far greater number,
viz. 30, have proved to be new to science, and of the genera two
have previously been known only from salt or brackish water.
Finally, one new genus, “rgasiloides, has been established, to
comprise three species allied to Hrgasilus Nordman.
The annexed table is intended to show the distribution in the
three lakes of the species here recorded, and, at the same time,
the number of species found in each of them. It will at once
appear from this table that Lake Tanganyika is by far the
richest in Copepoda, no less than 29 species having been recorded
from there, whereas a rather limited number of species is found
in the two other lakes, viz., in Nyasa 11, in Victoria Nyanza
only 7 species. This agrees pretty well with the results which
other authors have obtained, in regard to the richness and
specialisation of forms in that lake, as compared with the fauna
of the other African lakes.
In striking contrast hereto stands, however, the apparently
total absence in Tanganyika of Cladocera. Although I have
with the greatest care sought for forms of this group in the
numerous ‘samples from this lake, I have only succeeded in
finding in one of them a solitary specimen of a Moina, and
this specimen in all probability has only quite accidentally
been carried into the lake from some neighbouring stream.
a
68 PROF. G. O. SARS ON THE COPEPODA [Jan. 12,
‘able of Distribution.
| Europe
. : Jictoria| and other
s of Species. | Tanganyika. Nyasa. Vie
Names of Species | ganyika. Nyas INy aime.| aeig O°
| the world. |
u
|
| Diaptomus galeboides G. O. S...... 2 i e as
Ss mappremee) ro O} Se ca scbeco coc bee. sie
a stuhlmanni MWrazek ...... fe +
a Simplex Gan OspSseeeeeeeeee +
- cunningtoni G. O. S....... +. =
Schizopera mopimata G. O. 8. ......... +
re valiohiore C4, Os Soeno co seco +
i consimilis G. OL S.......... + a:
. uneulata GaN Si en +
ee minuticornis G. O. S....... | as
5 spinulosa G. O. S. .........) + |
a fimbriata G. O.S. fr 45 +
scalaris G. O.S. . +
| iiieor tats perplexus G. O. Ss. ae
| Cyclops leuckarti Claws Baader + + + +
| 5) CUMIN JER EAAY scoocaonsccacscece as ant =
E neglectus G. O. S. ............ + + = +
=A manele C2 Os Ss ccccocceondouss +
ss SMOG SHA aco cc. cos see catase! : + bs -
K atbenmatusy GON Sei eens +
| - varicans G. O. S. Me + ae a
| eexd cus) Gl Og See eee +
| cunningtoni G: O)7S) 7. +
s pachycomus G. O. S......... >
Ms semiserratus Gt. O. S.......... -
is levimargo G. O. S. +
a angustus G. O. S. + +
3 TALIS PINUS! Gy On Sewer eee +
a benllonsles Ee WL Sis ees condncee 25 =
x euacanthus GQ. 8S. ......... +
s gilenns G4 Qs Ss stecvecss cee cos +
3 oligarthrus G. O.S. ......... +
5 Gomes Ex OaiSo sccacecos 22
dubius G. O. S. Beat Bee +
| Ergasiloides megacheir G. (Oh SHODS +
* macrodactylus, GOS ¢ +
- brevimanus G. O. S. ... > +
JOREAISTIIG Soy“ seudadcoossy dongsooonsosecnces ae +
Dr. Cunnington also has noticed this peculiar feature of Lake
Tanganyika. In the other two lakes, on the other hand, a
number of well-marked species of that group occur, both true
limnetic and bottom-forms.
Of the six genera of Copepoda here recorded, the genus Cyclops:
is represented by far the greatest number of species, viz., in all
20; next to it comes the genus Schizopera with 8 species, the
genus Diaptomus with 5 species, the genus Hrgasiloides with
3 species, and finally the genera Lh yophilus and Hrgasilus, each
with a single species.
As to the distribution of the species, it ought to be noted that
the three takes have each their particular forms of Diaptomi not
found out of the respective lakes, one species occurring In
1909. | OF THE THIRD TANGANYIKA EXPEDITION. 69
Tanganyika, two in Nyasa, and two others in Victoria Nyanza.
Of the genus Schizopera a single species (S. fimbriata) seems to
be peculiar to Lake Nyasa; all the other seven species are
found in Tanganyika, and of these only one (S. consimilis) is
stated to occur also in one of the other lakes (Victoria Nyanza).
The remarkable genus Jlyophilus, as above stated, is only re-
presented by a single species from 'Tanganyika. Of the 20 species
of the genus Cyclops, 17 have been stated to occur in Tanganyika.
Two of these (C. lewckarti and C. neglectus) are common to
all three lakes; three species (C. varicans, C. angustus, and
C. agiloides) occur occasionally also in the other lakes, the
first two in Nyasa, the last in Victoria Nyanza. The remaining
12 species, on the other hand, seem to be endemic forms of
Lake Tanganyika, Of the three species which have not yet been
found in Tanganyika, one ((. emini) seems to be an endemic
form of Victoria Nyanza; the second (C. albidus) is a widely
distributed, almost cosmopolitan species, like C.. leuckarti; and the
third (C’, dubius) is as yet only known from a solitary specimen
found in Nyasa. The three species of the genus Hrgasiloides
occur all in Tanganyika, and only one of them (C. brevimanus) is
occasionally also found in Nyasa. In the latter lake, finally, a
still undetermined species of the genus Hrgasilus occurs.
Particular attention ought to be paid to the two remarkable
genera, Schizopera and Jlyophilus. Both these genera must
evidently be regarded as of marine origin, and the question thus
arises, how we shall explain the occurrence of species of these
genera in the purely freshwater lakes of Central Africa. The
most obvious inference appears to be the belief, that these species
are true “ relict” forms, that is to say, the remains of an ancient
marine fauna prevailing here at a time when the lakes formed part
of the Ocean; and indeed a supposition in favour of such a con-
clusion was advanced some time ago by Mr. J. E. 8. Moore,
who is of opinion that Lake Tanganyika might be the modified
remains of part of an ancient Jurassic Sea, and that its fauna
accordingly in some instances exhibits distinct traces of more
primitive (marine) characters. Recent investigation of this lake
tends, however, to disprove the supposition set forth by Mr. Moore
about the fauna of Lake Tanganyika, and to show that it is on
the contrary a highly specialised one, and does not exhibit any
true relation to marine forms. Jam myself also of opinion that
the theory of Mr. Moore about his so-called “ halolimnic ” (relict)
forms can scarcely be supported. But, how are we to explain
the presence in Lake Tanganyika of species of the two above-
mentioned genera? I think that we need not regard these as
true “relict” forms, although their marine origin seems to be
indisputable. We have in this case recourse to another ex-
planation, which may prove to be fully sufficient, namely, the
accidental transport by the aid of migratory aquatic birds. 'The
importance of such a transport for the distribution of small fresh-
water animals-has long been recognised. It is indeed easily
70 PROF. G. 0. SARS ON THE COPEPODA [Jan. 12,
understood, that not rarely it may happen that parcels of mud
adhering to the feet of such birds and containing germs
of small organisms, may be transported from one basin to
another for rather a long distance. Several of these germs, for
instance the resting ova of Rotatoria, Planaria, Bryozoa, and
small Crustacea, may be kept for years in a dried condition,
without losing their developing power, and of course will easily
develop when brought under favourable conditions. Even in
the case of animals, which do not produce such resting ova,
a successful transport in this way may be effected. A very
interesting discovery has recently been made by Prof. Birge and
Mr. Juday at the Laboratory of the Wisconsin Geol. and Nat.
Hist. Survey, in the case of a common species of the genus
Cyclops (C. bieuspidatus Claus)*. It has been stated by these
authors that this form, at certain periods of the season, is
subjected to a peculiar encysting process, small, still immature
specimens enveloping themselves with a rather firm cocoon of
muddy particles held together by some glutinous matter. It is
very easy to believe that such cocoons may be kept in a dry
condition for a long time without any damage to the enclosed
young Cyclops, and that consequently an accidental transport of
them by migratory birds may be as successful as that of true
resting ova. It also appears very probable that a similar en-
cysting process may be found to be present in other species of
this genus. In any case, the discovery of Prof. Birge and
Mr. Juday cannot fail to throw an unexpected light upon
certain difficult questions regarding the distribution of some
species of the genus Cyclops, and perhaps also of forms not
belonging to that genus.
The above remarks on the accidental transport of animals by
the aid of migratory birds, chiefly concerns true freshwater forms.
However, as the said birds not only visit freshwater lakes. but
also the coasts of the sea, it is evident that a transport of mud
from the sea-shores to freshwater lakes may occasionally take
place. In most cases, certainly, the germs contained in such
mud will not develop when brought into purely fresh water,
yet it is not impossible that the development of some few forms
may in reality be effected under such circumstances. This
may be assumed to be the case with marine animals that have
accustomed themselves to live in more or less brackish water,
and this is precisely the case with the type species of both the
two above-mentioned genera. One of them, /lyophilus flexibilis,
has even been stated by Prof. Lilljeborg to occur occasionally
also in purely fresh water, having been found, besides in the
Baltic, also in the neighbouring Lake Melaren, and this fact
clearly proves that such brackish water animals in reality possess
the power of accustoming themselves to living in fresh water.
It has been stated above, that no less than eight different
* “ A summer resting-stage in the development of Cyclops bicuspidatus Cls.”
1909. | OF THE THIRD TANGANYIKA EXPEDITION, vt
species of the genus Schizopera have been found in the Central
African lakes, seven of them occurring in Lake Tanganyika. On
a closer examination, it has been proved that none of these
species is identical with the type-species, S. longicauda, ex-
hibiting as they do some well-marked differences of apparently
specific value. Of course, it would be quite unreasonable to
assume that all these species have been transported to the lake in
the above-mentioned manner. In my opinion these species have
developed independently in the lake from a single ancestral form
(perhaps S. longicauda), which in some remote time has found
its way to the lake in the above-mentioned manner, and which,
owing to the altered condition of life and isolation, has gradually
changed its characters and undergone a divergent development
into several varieties. These varieties in their turn have at last
attained the character of distinct species.
In a similar manner the African species, //yophilus perplexus,
may have been originally derived from the type species, /. flexi-
bilis, though the alteration of characters, which has taken place,
is great enough to distinguish it as a well-marked species. The
occasional occurrence of two species of the genus Schizopera in
the two other African lakes, in all probability is due to an acci-
dental transport from the neighbouring Lake Tanganyika, and
the same may also be the case with some other Copepod-species
stated to occur both in that lake and in one or other of the two
remaining lakes. It may be observed here, that Lake Tanganyika
in its general physical characters differs from the other two
lakes. It is extremely long and narrow, in some places exhibit-
ing very considerable depths, and we have historical evidence to
prove that its water was formerly somewhat brackish in character.
Nyasa, it is true, while not so long, is also deep and relatively
narrow, but Victoria Nyanza is very broad and quite shallow.
The above-mentioned and other physical and perhaps also bio-
logical peculiarities of Lake Tanganyika seem to have favoured,
during a long period of isolation, a divergent development of
certain species, and this development in some cases has preceeded
to such an extent as to produce even new generic types, all of
which, however, have conserved the stamp of their phylogenetic
relation to other forms occurring in the same lake or otherwise.
Such highly specialised forms, representing particular genera,
have been recorded among the Brachyura by Dr. Cunnington,
and among the Macrura by Dr. Calman. Also among the Cope-
poda a new generic type has been found, viz., 41 ‘gasiloides, with
three well-marked species. The phylogenetic relation of this
genus to the genus Hrgasilus is very obvious, the generic difference
chiefly consisting in a somewhat retrograde transformation of
the posterior part of the body.
A development, in comparatively recent times, of new species
and even genera in isolated basins, is by no means a unique
feature exclusively peculiar to Lake Tanganyika. We know of
similar cases also from other parts of the world. Thus, it is well
re
12
known that Lake Baikal in Siberia distinguishes itself in a
quite remarkable manner from the other Asiatic lakes by its
astonishing richness in various species of Gammarid Amphipoda.
It is impossible to explain this peculiarity of Lake Baikal without
assuming that ‘a divergent development from one or a few
ancestral forms has taken place here.
we meet with a quite similar case, not only in the Amphi-
poda, but also in other groups of Crustacea, viz. Mysidee, Cumacea,
Polyphemide, as is shown by the present author in his several
PROF. G. 0. SARS ON THE COPEPODA [Jan. 12,
papers on the Carcinological Fauna of that interesting basin.
Fic.
Fig.
Wig.
18.
19.
20.
PAE
22.
2:
24.
5. GY
g@. Zo.
26.
27.
28.
29.
30.
21.
29
EXPLANATION OF THE PLATES.
PrarE VI.
Diaptomus galeboides G. O. Sars.
. Adult, ovigerous female, dorsal view.
. Same, viewed from left side.
Urosome together with part’ of metasome, more highly magnified
and viewed from the dorsal face.
. Right lateral corner of last segment of metasome, still more highly
c t= ? > J
magnified.
. Leg of last pair.
. Adult male, dorsal view.
. Terminal section of right anterior antenna.
. Last pair of legs.
Prare VII.
Diaptonmus mivtus GO. Sars.
. Adult female, dorsal view.
. Same, viewed from left side.
. Posterior part of metasome and genital segment, dorsal view.
2. Leg of last pair.
. Adult male, dorsal view.
. Middle and terminal sections of right anterior antenna.
. End of terminal part of same antenna, more highly magnified.
. Last pair of legs.
Projection of the 2nd basal jomt of right leg, more highly magnified.
Prate VIII.
Diaptomus stuhimanni Mrazek.
Adult ovigerous female, dorsal view.
Same, viewed from left side.
Left lateral part of last segment of metasome.
Leg of last pair.
Terminal part of right anterior antenna in male.
. Last pair of legs of same.
Outer ramus of left leg, viewed from the anterior face, more highly mag-
nified.
Diaptomus simplex G. O. Sars.
Adult ovigerous female, dorsal view.
Same, viewed from left side.
Urosome together with posterior part of metasome, dorsal view.
Right lateral part of last segment of metasome, somewhat more maguificd,
Leg of last pair.
Outer three joints of right anterior antenna of male.
Last pair of legs of same.
Outer ramus of left leg, viewed from the anterior face, and more highly
magnified.
In the Caspian Sea, too,
1909.]
Fig.
Fig. /
Fic.
Fig.
Fig.
34.
35.
36.
aly
38.
39.
40.
4),
42,
OF THE THIRD TANGANYIKA EXPEDITION. 73
Prate IX.
Diaptomus cunningtoni G. O. Sars.
Adult female, dorsal view.
Another female, with attached spermatophore. viewed from left side.
Urosome together with posterior part of metasome, dorsal view.
Left lateral part of last segment of metasome, with adjoining part of genital
segment, dorsal view.
Same part viewed from the exterior face.
Last pair of legs.
Adult male, dorsal view.
Terminal part of right anterior antenna of same.
Last pair of legs,
Outer ramus of left leg, viewed from the anterior face, and more highly
magnified.
PLATE X.
Schizopera inopinata G. O. Sars.
. Adult ovigerous female, dorsal view.
|. Same, viewed from left side.
tostrum together with right anterior antenna, dorsal view.
Right posterior antenna.
. Outer ramus of same, highly magnified.
. Mandible with palp.
. Posterior maxilliped.
50. Leg of 1st pair.
. Leg of 2nd pair.
. Leg of 3rd pair.
53. Leg of 4th pair.
54. Last pair of legs.
. Extremity of urosome, with the caudal rami, dorsal view.
56. Anterior antenna of male.
. Inner ramus of a leg of 2nd pair of same.
. Leg of last pair of same.
Prate XI.
Schizopera validior G. O. Sars.
Adult ovigerous female, dorsal view.
Anterior antenna.
Leg of 1st pair.
. Leg of last pair.
. Right caudal ramus.
Schizopera consimilis G. O. Sars.
Adult ovigerous female, dorsal view.
5. Leg of 1st pair.
Leg of last pair.
. Right caudal ramus.
Schizopera ungulata G. O. Sars.
Adult female, dorsal view.
Leg of 1st pair.
. Leg of last pair.
. Left caudal ramus.
Priate XII.
Schizopera minuticornis G. O. Sars.
Adult female, dorsal view. ;
Rostrum together with right anterior antenna, dorsal view.
. Leg of 1st pair.
Leg of 2nd pair.
Leg of last pair.
. Left caudal ramus, with adjoining part of urosome, dorsal view.
74 PROF. G. 0. SARS ON THE COPEPODA [Jan. 12,
Schizopera spinulosa G. O. Sars.
Fig. 78. Adult female, dorsal view.
79. Leg of Ist pair.
80. Leg of last pair.
81. Lett caudal ramus, with adjoining part of urosome, dorsal view.
Schizopera fimbriata G. O: Sars.
Fig. 82. Female (not fully grown), dorsal view.
83. Leg of Ist pair.
84. Leg of 4th pair.
85. Leg of last pair.
86. Left caudal ramus, with adjoining part of urosome, dorsal view.
PuatE XIII.
Schizopera scalaris G. O. Sars.
7. Adult male, dorsal view.
8. Anterior antenna.
9. Leg of Ist pair.
0. Leg of 2nd pair.
91. Leg of last pair.
92. Extremity of urosome, with right caudal ramus, dorsal view.
Fie.
Tlyophilus perplexus G. O. Sars.
Fig. 98. Adult female, dorsal view.
94. Same, viewed from left side.
95. Rostrum, together with left anterior antenna, dorsal view
96. Right posterior antenna.
97. Mandible with palp.
98. Manilla.
99. Anterior mavilliped.
100. Posterior maxilliped.
101. Leg of Ist pair.
102. Leg of 8rd pair.
103. Leg of 4th pair.
104. Leg of last pair.
=
io
Prate XIV.
Cyclops leuckarti Claus.
Fig. 105. Adult female, dorsal view.
106. Terminal joint of inner ramus of a leg of 4th pair.
107. Leg of last pair.
Cyclops emini Mrazek.
Fig. 108. Adult ovigerous female, dorsal view.
109. Posterior antenna.
110. Inner ramus of a leg of 4th pair.
111. Leg of last pair.
112. Extremity of urosome, with the caudal rami, dorsal view.
Cyclops neglectus G. O. Sars.
Fig. 113. Adult ovigerous female. dorsal view.
114. Posterior antenna.
115. Inner ramus of a leg of 4th pair.
116. Leg of last pair.
117. Extremity of urosome, with the caudal rami, dorsal view.
1909.]
Fig,
Fig.
Fig.
118.
119.
120.
121.
122.
123.
124.
125.
126.
127.
. 128.
129.
130.
131.
132.
133.
134.
135.
136.
137.
138.
@. 139.
140.
ig. 141.
142.
ig. 143.
144.
145.
146.
147.
148.
149.
150.
151.
152.
153.
154.
155.
156.
OF THE THIRD TANGANYIKA EXPEDITION.
PratE XV.
Cyclops tenellus G. O. Sars.
Adult female, dorsal view.
Anterior antenna.
Posterior antenna.
The two maxillipeds on left side.
Leg of Ist pair.
Leg of 2nd pair.
Outer ramus of a leg of 3rd pair.
Lege of 4th pair.
Leg of last paiv.
Extremity of urosome with the caudal rami, dorsal view.
Cyclops albidus (Jurine).
Adult male, dorsal view.
Inner ramus of a lee of 4th pair.
75
Right half of the last pedigerous segment and of the genital segment
viewed from the ventral face, exhibiting the corresponding leg of last
pair and genital lamella, as also an enclosed spermatophore.
PraAtTE XVI.
Cyclops attenuatus G. O. Sars.
Adult ovigerous female, dorsal view.
Anterior antenna.
Posterior antenna.
Leg of 1st pair.
Lee of 3rd pair.
Lee of 4th pair.
Lateral part of last pedigerous segment, with the corresponding
rudimentary leg.
Extremity of urosome, with the caudal rami, dorsal view.
Cyclops varicans G. O. Sars.
Adult female, dorsal view.
Extremity of urosome, with the caudal rami.
Cyclops exiguus G. O. Sars.
Adult female, dorsal view.
Extremity of urosome, with the caudal rami.
PratE XVII.
Cyclops cunningtoni G. O. Sars.
Adult ovigerous female, dorsal view.
Anterior antenna.
Posterior antenna.
Leg of 1st pair.
Leg of 3rd pair.
Leg of 4th pair.
Lateral part of last pedigerous segment, with the corresponding
rudimentary leg.
Left caudal ramus, with adjoining part of urosome, dorsal view.
Cyclops pachycomus G. O. Sars.
Adult female, dorsal view. °
Anterior antenna.
Posterior antenna.
Leg of 4th pair.
Lateral part of last pedigerous segment, with the corresponding
rudimentary leg.
Extremity of urosome, with the caudal rami, dorsal view.
76 PROF. G. 0. SARS ON THE COPEPODA [Jan. 12
PratEe XVIIT.
Cyclops semiserratus G. O. Sars.
Fig. 157. Adult ovigerous female, dorsal view.
158. Anterior antenna.
159. Posterior antenna.
160. Mandible with rudimentary palp.
161. Mavwilla.
162. Anterior maxilliped.
163. Posterior maxilliped.
164. Leg of 1st pair.
165. Leg of 8rd pair.
166. Leg of 4th pair.
167. Lateral part of last pedigerous segment, with adjoining part of genital
segment, exhibiting the corresponding leg and seta, ventral view.
168. Leg of last pair isolated and more highly magnified.
169. Extremity of urosome, with the caudal rami, dorsal view.
PrarEe XIX.
Cyclops levimargo G. O. Sars.
Fig. 170. Adult ovigerous female, dorsal view.
171. Anterior antenna.
172. Posterior antenna.
173. Leg of 4th pair.
174. Leg of last pair.
175. Right caudal ramus, with adjoining part of urosome, dorsal view.
Cyclops angustus G. O. Sars.
Fig. 176. Adult ovigerous female, dorsal view.
177. Outer part of an anterior antenna.
178. Posterior antenna.
179. Leg of last pair.
180. Right caudal ramus, with adjoining part of urosome, dorsal view.
Cyclops rarispinus G. O. Sars.
Fig. 181. Adult ovigerous female, dorsal view.
182. Left caudal ramus, with adjoining part of urosome.
PratE XX.
Cyclops agiloides G. O. Sars.
Fig. 183. Adult female, dorsal view.
184. Outer part of an anterior antenna.
185. Posterior antenna.
186. Leg of 4th pair.
187. Leg of last pair.
188. Lett caudal ramus, with adjoining part of urosome, dorsal view.
Cyclops euacanthus G. O. Sars.
Fig. 189. Adult ovigerous female, dorsal view.
190. Extremity of an anterior antenna.
191. Leg of 4th pair.
192. Right caudal ramus, with adjoining part of last segment, dorsal view.
Cyclops ciliatus G. O. Sars.
Fig. 193. Adult female, dorsal view.
194. Left caudal ramus, with adjoining part of urosome, dorsal view.
1909.
Fig.
Fig.
Fig.
208.
209.
210.
PALE
212
—
213.
214.
215.
216.
217.
218.
219.
220.
221.
999
aaa:
eis
9 bo
He Co
220.
226.
227.
228.
229
—_
OF THE THIRD TANGANYIKA EXPEDITION. 77
PratE XXI.
Cyclops oligarthrus G. O. Sars.
. Adult ovigerous female, dorsal view.
. Anterior antenna.
. Posterior antenna.
. Leg of 1st pair.
. Leg of 3rd pair.
. Inner ramus of a leg of 4th pair.
. Lateral part of last pedigerous segment, with the corresponding leg.
. Right caudal ramus, with adjoining part of anal segment, dorsal view.
Cyclops compactus G. O. Sars.
. Adult female, dorsal view.
. Lateral parts of the last two pedigerous segments and of the genital
segment, exhibiting the corresponding leg of last pair, and the short
seta, dorsal view.
Cyclops dubius G. O. Sars.
5. Adult female, dorsal view.
. Posterior part of metasome and adjoining part of genital seement, viewed
from the dorsal face, and exhibiting the peculiar ciliation of the last
pedigerous segment, and the projecting legs of last pair.
. Left caudal ramus, with adjoining part of urosome, dorsal view.
Prats XXII.
Ergasiloides megacheir G. O. Sars.
Female in last free stage, viewed from dorsal face.
Same, viewed from left side.
Anterior antenna.
Posterior prehensile antenna.
Oral area, viewed from the ventral face, and exhibiting the oral parts
an situ.
Mandible highly magnified.
Manilla.
Rudimentary maxilliped.
Leg of Ist pair.
Leg of 3rd pair.
Leg of 4th pair.
Rudimentary leg of last pair, together with part of the likewise rudi-
mentary corresponding segment.
Adult male, dorsal view.
Posterior antenna of same.
Maxilliped of same.
Prate XXIII.
Ergasiloides macrodactylus G. O. Sars.
. Female in last free stage, dorsal view.
. Posterior prehensile antenna.
Ergasiloides brevimanus G. O. Sars.
Female in last free stage, dorsal view.
Posterior prehensile antenna.
Leg of 1st pair.
Leg of 3rd pair.
. Leg of 4th pair.
230.
231.
Urosome, with the caudal rami, dorsal view.
Male, viewed from the dorsal face.
78 MR, T. GOODEY ON THE [Jan. 12,
3. A further Note on the Gonadial Grooves of a Medusa,
Aurelia aurita. By T. Goopry, B.Se., University Scholar,
Zoological Laboratory, University of Birmingham.*
[Received November 19, 1908. }
(Plate XXLV.+)
In a short paper read before this Society in February last +
the occurrence of these grooves was noted and their structure
and relations figured and described. The present paper is the
outcome of observations which were suggested in the earlier
communication and which prove that the grooves actually function
as gonoduets. For the purpose, however, of making this paper
complete in itself, it has been thought desirable to give a brief
résumé of what the former paper dealt with.
Tt was pointed out that the grooves are four narrow channels,
best seen in a subumbrella aspect of the medusa, lying in the
interradial axes. They have about the same diameter as the
origins of the ordinary gastrov aseular canals, and are confined to
the ventral walls or floors of the main passages from the central
gastric cavity to the gastric pouches, and to the floors of the
gastric pouches themselves. ‘They terminate in slightly funnel-
shaped expansions at about the centre of each pouch. The
boundaries of each groove are formed by two parallel ridges of
epithelium which are raised up so as to form a channel between
them. It was suggested that, functionally, the grooves serve as
channels along which the sex-cells pass on their way to the
exterior after having been shed into the gastric pouches at the
time of spawning ; that they were, in short, incipient gonoducts
eut off from the ‘general archenterie cavity or celenteron.
It seemed desirable to ascertain whether these views were
correct by the examination of specimens taken at the period of
sexual maturity.
In order to undertake this investigation I spent several weeks
in the laboratory of the Marine Biological Station at Plymouth
during the month of August, and there collected the necessary
material. The specimens were all taken from the open sea and
were of the type with very thick mesoglea; those worked on
in the University laboratory for the first paper were estuarine
specimens and were compar atively thin and discoidal. A large
number of meduse, about two hundred in all, were examined with
the view of finding sex-cells in the gonatlial grooves. Of these
only a few, about tw elve, possessed grooves In which any obvious
contents could be detected. Apparently the reason why | compara-
tively few specimens exhibited grooves with eontents, was that
* Communicated by Prof. T. W. Brrpes, Se.D., F.R.S., F.Z.S,.
+ For explanation of the Plate see p. 81.
* 'T. Goodey, P. Z.S. 1908, p. 55.
PS 1590S P1 Sx:
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West, Newman lth
IA AURITA.
di
GROOVES OF AURE
GONADIAL
T.G. del.
1909. ] GONADIAL GROOVES OF A MEDUSA. is
out of the large number taken only one here and there was
at the exact stage when the sex-cells were being extruded ;
and, further, it appears that the whole of any particular gonad-loop
is not rive at one and the same time, so that only a portion of it
may be shed at a time and so perhaps be found in the gonadial
groove.
After taking these facts into consideration, it is gratifying to
have been able to find some specimens in which the grooves
showed unmistakable indications of the presence of sex-cells.
In one or two cases the small masses of material were with-
drawn from the grooves by means of a fine pipette and then
examined under the microscope. By this means both eggs and
masses of spermatozoa were recognised. However, it was not
possible to figure these structures accurately under these
conditions ; and although it was proof that the grooves actually
serve as gonoducts, yet it was deemed necessary to obtain figures
of the sex-cells in the groove. For this purpose, the remainder of
the specimens which showed grooves with contents were dissected
and the necessary portions were removed entire. These latter
were stained in bulk in borax-carmine for forty-eight hours,
dehydrated, and embedded in paraffin-wax. Sections were cut at
right angles to the interradial axes along which the grooves lie.
In the preparation of the material for sectionizing, there was a
large amount of contraction owing to the great thickness of the
mesogloal tissue, and for this reason the lining epithelium in
many places became broken up and displaced from its original
position. In spite of this, however, the grooves have retained
their position and proportions fairly well, and it has been possible
to obtain figures of the sex-cells in sitv.
The first figure (Pl. XXIV. fig. 1) shows a gonadial groove the
lining epithelium of which (e) has been fragmented somewhat.
A mass of nucleated tissue is shown lying within the limits of
the groove. On the examination of this with high power
objectives (fig. 2) it was seen to consist of spermatozoa. Along
with the latter there are other large cells which are in
various stages of disintegration and are apparently the nurse
or nutrient cells (Aders*). The spermatozoa in the middle
of the figure can be easily made out, each having a deeply stained
nuclear head. In the case of a few spermatozoa, the clear,
almost globulax middle-piece can be distinguished. Flagella are
also very numerous in the section. From the densely aggre-
gated nature of the two masses of male cells at the top and bottom
of the figure, and also from their relations with the nutrient
cells, it would seem as though spermatogenesis, or at least some
of the final stages of the process, may take place within the limits
of the groove.
Fig. 3 shows a mass of eggs lying in a groove. Unfortunately,
the piece of material from which this section was cut turned
* Zeits. f. wiss. Zool. Ixxiv. 1903. p. 95.
80 ON THE GONADIAL GROOVES OF A MEDUSA. [Jan. 12,
out to be very difficult to manipulate in the preparation for
sectionizing. There was a very great amount of contraction of
the mesogloal tissue, so that the sections are considerably distorted.
However, the figure shows the limits of the groove fairly well
with its contained mass of eggs.
The above account affords pretty conclusive proof, I think, of
the suggestion that these gonadial grooves function as gonoducts,
and that it is only sriintn their limits that the sex-cells pass on
their way to the exterior. Also out of ali the specimens
examined, only comparatively few, as mentioned before, showed
anything in the space between the gastric pouch and the central
gastric cavity, and in all of these the substance lay in the grooves
themselves.
Thus Awrelia may be looked upon as possessing ccelomic or
archenteric derivatives of the nature of primitive gonoducts,
standing alone in this respect among the Ccelenterata so far as
we know at present, for the origin of the genital ducts of
Ctenoplana korotneffii is not yet known.
It is of interest to note that other observations along different
lines point to the grooves being connected in some way with the
gonads. From an examination of a series of specimens of
different stages in development, including ephyre and small but
complete smell of about one inch in diameter, the gradual
differentiation of the gastro-vascular system can be traced. In
those specimens which show the completely developed gastro-
vascular canals, there are slight indications of gonadial grooves in
the floors of the almost fully developed gastric pouches. The latter
are formed by outgrowths of the central or general gastric cavity
along the interradii and by inpushings of this same gastric
cavity along the pervadii. By the last mentioned process the
main passages of connection between the central gastric cavity
and the gastric pouches are established. While this process is
going on, “the gonadial grooves appear as two parallel ridges in the
ventral wall of each main passage and gastric pouch. From this
it will be seen that the gonadial grooves are of later appearance
than the ordinary radial “canals, Their complete differentiation
is coincident with the final stage in the growth of the gastric
pouches, and incidentally they approach completion just prior to
the appearance of the gonads.
From this we may safely infer that the gonadial grooves are
not concerned with a nutrient function, but rather that they are
formed in readiness to discharge their function in connection
with the gonads.
‘Thr ough the kindness of the officials at the British Museum, I
was able to examine a number of Acraspedote medusze in
January last. As a result of this I am able to record the
presence of the gonadial grooves in another species of the genus
Aurelia, viz. oman ried. and in three varieties of Aurelia
aurita, viz., A. aurita var. japonica; A. aurita var. cruciata ;
and A. aurita var. colpota.
1909. ] ON THE TUBERCULIN TEST IN MONKEYS, 81
Pilema octopus, Pelagia phosphora, and three specimens of
‘Cassiopea were also carefully examined, but no trace of gonadial
grooves could be found in any of them.
In conclusion I desire to tender my warmest thanks to Lord
Avebury for his kindness in nominating me to the use of a. table
-at the Plymouth laboratory during the month of August, and also
to the laboratory officials for the excellent facilities which were
-afforded me for the collection and examination of specimens during
my stay there.
EXPLANATION OF PLATE XXIV.
Gonadial grooves of Aurelia aurita.
All the figures are camera lucida drawings. Fig. 2 is with Leitz —, oil-immersion
; 2 5 te) 5 2
“objective and No. 4 eyepiece.
‘Fig. 1 (X40). Section passing through groove, at a point just within the gastric
pouch. Thick buttressing mesogloea on either side of the groove. Mass
of male cells lying within the groove.
| Fig. 2 (1050). Portion of section in fig. 1 showing spermatozoa and nutrient
cells in the groove.
Fig. 3 (X46). Section passing through main passage to a gastric pouch, showing
gonadial groove in the floor in an almost median position bounded on either
side by endodermal epithelial folds. Between the latter is a mass of eggs.
REFERENCE LETTERS.
b.w, body-wall or mesoglea; d.n.c, disintegrating nutrient cells; e, epithelium
lining gonadial groove and main passage; g.g, gonadial groove; g.p, gastric pouch;
n.c, nutrient cells; 0, eggs; s.p, spermatozoa.
4, The Tuberculin Test in Monkeys: with Notes on the
Temperature of Mammals. By Arraur Erwin Brown,
D.Sc., C.M.Z.S., Secretary of the Zoological Society of
Philadelphia.
[Received November 25, 1908. |
(Text-figures 1—4.)
For many years previous to the introduction of a rigorous
‘quarantine system for incoming monkeys, and of exact measures
for the detection of tuberculosis in the Zoological Gardens at
Philadelphia, it is believed that at any given time from one-fifth
to one-fourth of the monkeys in the collection were tuberculous.
Indeed, there have been at least two periods when, under what
appeared to be an infection of especial virulence, the death-rate
-exceeded this average.
A more or less similar experience has been that of the older
Zoological Gardeus in Europe and America,
In March 1905, upon the opening of the Laboratory of Patho-
logy in the Gardens at Philadelphia, a Series of observatious was
-begun by Dr C. Y. White, at that time the Society’s pathologist,
Proc. Zoou. Soc.—1909, No. VI. 6
82 DR. A. E, BROWN ON THE | Jan. 12,
and myself, having in view the possible control of this disease.
Tt was known that many of the monkeys coming to the Gardens
from the hands of dealers were already infected, and that even in
advanced stages they are often of healthy appearance and seldom
give reliable external indications of the disease.
The old stock of monkeys had been undergoing a period of
acute infection, and without exception they were removed from
the Monkey House and for the most part used for experimental
purposes, the building itself being thoroughly cleaned and dis-
infected with formaldehyde vapour. Since then every monkey
reaching the Gardens has been placed in a quarantine room, and
only those that successfully passed the test of sub-cutaneous
injection of tuberculin have been sent to the Monkey House for
exhibition. The use of tuberculin has thus been solely for a
diagnostic purpose.
Many difficulties were encountered in the early stages of the
work, mainly due to the irregularities of temperature in monkeys,
and to our initial ignorance of what should be regarded as normal
in healthy animals, as well as of the kinds of reaction to be
expected in tuberculous cases.
These obstacles were for the most part overcome through
careful study of the conditions shown on autopsy in a large
number of animals, in connection with the corresponding tempera-
ture charts, and it is now felt that the method and the results
have reached a large measure of certainty.
The individual variations in temperature are marked, and it is
frequently necessary to take a record (rectal in all cases) daily for
several weeks before it settles down to approximate regularity,
which in our experience is about 101° or 102° Fahr. in the mid-
afternoon. This is slightly higher than the 38° Cent. (100-4° F.)
assumed as a normal by Simpson and Galbraith*, but it is
probable that the difference may be accounted for by the fact
noted by us and also shown by those authors, that excitement and
muscular effort definitely raises temperature in these animals 7.
The monkeys used in their observations were handled for a much
longer period than ours, and they were able to select those that
became relatively indifferent to the usage and restraint involved.
Although we have accepted the above standard as normal, cases
have occurred where monkeys have shown temperatures never
below 104° at the daily maximum, and are still living and to all
appearance in good health.
An important fact was early determined by a sequence of
* Trans. Royal Soc. of Edinburgh, xlv. pt. 1, p. 98 (1906).
+ An interesting instance is afforded by a lot of five monkeys subjected to the
test a few weeks ago. Temperatures were taken daily for two weeks before injection
and all van with unusual regularity. On September 25th, near the end of the period,
each chart showed a sharp elevation of from “4° to 12° F. It was found upon
inquiry that the attendant who usually took the temperatures was absent on that
day, and the duty fell to another man who had experience in the work but was less
gentle in handling the animals, and who, moreover, was quite unknown to this
particular lot.
1909. | TUBERCULIN TES! IN MONKEYS. 83
records taken at intervals of four hours during several days, in the
existence of a marked daily rhythm or range in the case of every
species of monkey examined, the high point being reached toward
3 p.M., about which time a steady fall sets in amounting to three
or four degrees by 3 A.M.; from then it rises again to the after-
noon maximum. This daily range is shown in Chart A (text-
fig. 1), and is in substantial agreement with the results reached
by Simpson and Galbraith. It should be mentioned that few
charts are as symmetrical as the one selected.
Text-fig. 1.
Chart A.
a)
[ i zal
| a -
- alt
TIME 7AMINA.MI3 P.MI7 PR MJB MJ3 A.Mj7 AMI A.M)3 PM.|7 P.M.j1LP.M.}3 A.Mj7A.M.|11A.M13 PM.
Cercopithecus pygerythrus $ adult.
Temperature chart showing normal daily curve.
In the beginning temperatures were taken for only twenty-four
hours after injection, but it was found that reactions are some-
times delayed until the second day, and an essential gain was made
in extending the observation period to forty-eight hours.
The course of procedure now followed in the tests is that when
the daily temperature of the individual monkey has settled down
to a somewhat regular course, an injection is made under the
loose skin of the lumbar region of from # to 2 milligrams of Koch’s
original tuberculin. This dose is only exceeded in case of ex-
ceptionally large animals, such as anubis or chacma_ baboons,
which have received as much as 3 mgs. Injection is made in the
forenoon, in order that a rise of temperature concurrent with the
upward tendency culminating about 3 P.M. may be noted. Tem-
eratures are then taken at intervals of four hours for two full
days. The hours we have selected as best indicating on the chart
the daily course, are 7, 11, and 3.
With healthy monkeys we have observed no change in the
degree or course of the daily rhythm, following injection, nor ill
results of any kind.
6*
84 DR. A. E. BROWN ON THE (Jan. 12,
In tuberculous subjects the usual result is a general rise of
temperature within the first twelve-hour period, though in a few
cases the rise has not appeared until the next day, or in many
cases a complete destruction of the daily curve ensues, with or
without a general rise. This last condition clearly represents the
balance reached in the struggle between two opposing tendencies,
the upward one due to tuberculin reaction and the downward one
which is the normal nightly course, and in meaning is equivalent
to a definite rise. Chart B (text-fig. 2) is of this character. In
this case, a young rhesus monkey, the temperature before injec-
tion ranged from 102°6° to 104-4°.
Text-fig. 2.
Chart B.
FFIME [TAM |TAM|SP.M.7 PM [I PMSAM|ZAM[TAM|SP.M|7PMINPM|SAM|7AMIITAM|S PA,
Macacus rhesus § juv.
Temperature chart showing tuberculous reaction destroying daily curve,
with no general rise.
It may be noted that in every case showing reaction of either
type, autopsy has shown tuberculous lesions *.
The charts most difficult of interpretation, however, in the
study of which quite a number of healthy monkeys were sacrificed,
are those not infrequent ones in which there is no definite rise in
temperature, and no complete destruction of the night drop, but
only more or less of a failure to properly complete it. These cases
are now held over for retesting after an interval of six or eight
weeks, and occasionally the second test gives a positive reaction.
It has not infrequently happened that monkeys have been retested
as many as three times before a final conclusion was reached.
Such charts call for nice discrimination, and, even with the most
expert, in them must always le a possibility of error.
In a few very advanced stages of general tuberculosis there was
no reaction rise, and the daily curve was completed, so that a
serious certainty of error would arise were it not for the fact that
* J am not prepared at this time to suggest an explanation of the difference
between our results and those announced by the Royal Commission on Tuberculosis
(Second Interim Report, p. 44, London, 1907). If anything has been certain in our
experience, it is that in not a single case of definite reaction has autopsy failed to
show tuberculosis.
1909. ] TUBERCULIN TEST IN MONKEYS.
85:
in every such case that we have observed, the whole temperature
after injection dropped conspicuously and death speedily followed.
Chart C (text-fig. 3) represents such a case.
Text-fig. 3.
Chart C.
JL
|
at
TIME |3 PMJ7P.MJIIPM13 AM)7AMII-A.M/3 PMJ7 P.M.}11P.M.)3 A.M17A.MJIA.M.|3 RM.)
Cercopithecus sabeus @ adult.
Temperature chart showing marked post-injection fall m advanced tuberculosis.
These advanced subjects, the only ones in which positive re-
actions have not been given in presence of tuberculosis, do not
impair the practical value of the test, for they are quickly
eliminated by death.
In one respect, mainly of pathological interest, much remains
to be determined, in the proportion between the degree of re-
action and the extent of the tuberculous lesions.
So far, we have
found an apparent lack of any such constant relation except as
noted in very advanced cases, in which, under the large doses of
tuberculin used, temperature has made a marked drop.
The necessity for an inflexible practice in the interpretation of
86 DR. A. E. BROWN ON THE [Jan. 12
charts cannot be too strongly stated. It has been followed
relentlessly throughout our work, and every doubtful case was in
the end decided against the monkey. In every case when an
animal was condemned, it was etherized and brought to the post-
mortem table.
A thoroughly complete system of prevention would perhaps
require the retesting of each monkey in the exhibition series at
least once a year, but this is no small under taking in an extensive
collection, and we have restricted the practice to those concerning
which suspicion arises based on ill-health or other indications.
These are immediately removed to the Laboratory and reinjected.
If the reaction is bad, the monkeys occupying the same cage are
at once taken to the Laboratory and put through the same course.
The infected cage as well as the adjoining ones are then thoroughly
disinfected,
Quite recently I have made a study of the history of every
monkey and lemur that has been tested since March 1905, in the
light of the records now in the Laboratory, consisting of two
hundred and sixty-seven post-injection temperature charts, and
complete post- -mortem records of nearly one hundred cases.
Up to November 1908, the test has been applied to one hundred
and sixty-three newly arrived monkeys belonging to Presbytis,
Cercopithecus, Cercocebus, Macacus, Cynopithecus, Papio, Ateles,
Cebus, Chrysothrix, Callithrix, and including a few of the genus
Lemur. Of these, one hundred and four passed the test and were
sent to the Monkey House. Of those not passed, twenty-five
were promptly condemned on their reactions and showed tuber-
culosis on autopsy. The remainder either died from other causes
while in quarantine, or were etherized to learn the meaning of
dubious charts, during the early and experimental stages of the
work,
Of the one hundred and four originally passed, ten have
developed tuberculosis during the three and a half years that have
elapsed, Three of these died in the exhibition cages, giving no
sign of ill-health. The others were detected and returned to the
Laborator y, Where they were either etherized or secluded. Seven
of these cases are traceable with reasonable certainty to two
infections from monkeys which were passed early in the tests on
twenty-four hour records, whose charts would be considered
doubtful in the ight of later experience. The three remaining
ones were also passed early in the tests, two of them on vather
uncertain charts, that of the third beimg of good quality though
somewhat irregular according to our pr esent standards. This
monkey, a male Cercopithecus ruber, tested in November 1906,
died six weeks later from nephritis and fatty degeneration of the
liver, and showed early miliary tuberculosis of the intestinal sub-
mucosa. With this exception, no monkey passed since February
1906 has as yet shown signs of tuberculosis, and more thana year
has elapsed since the last one exhibiting it was removed from the
Monkey House on October 16th, 1907.
1909. | TUBERCULIN TEST IN MONKEYS. 87
The history of this case,a male Cercocebus fuliginosus (Number
59), is interesting, not only because it was one of the first lot
tested, but as an example of the need for unceasing vigilance. It
reached the Gardens on March 29th, 1905, and had a temperature
taken almost daily until June 28th, the range being from 100-6?
to 103°8°. On the last date it was injected and gave a twenty-
four hour temperature which appears good, but not being fully
understood the animal was held over and reinjected on August 8th,
at which time it was passed, again on a twenty-four hour record.
In January 1906 it seemed unwell and was returned to the
Laboratory for retesting, on which occasion the forty-eight hour
chart was good, with trifling irregularities. It was kept under
observation until March 6th, when it was returned to the Monkey
House. Later in that month three others of the same species,
originally passed in December 1905, were removed for retesting
and found to be tuberculous, one of them (No. 62) having been the
source of infection, as is indicated by present reading of the charts.
Number 59 remained on exhibition in apparently good health for
nineteen months, until October 16th, 1907, when it was removed to
the Laboratory, dying three days later with general tuberculosis.
After March 1906 this animal was not exposed to infection from
other monkeys, and it is probable that it contracted the disease at
that time from its three cage-mates, which were found to be
tuberculous several weeks after its return.
Text-fig. 4,
Chart D.
Ba
(ea
ams em|7em |i PM.]3 AM
Lemur varius 2 adult.
Non-tuberculous temperature charts. Unbroken curve taken April 9th—-11th, 1907;
broken curve taken July 1st-3rd, 1907.
Progress of the disease is usually much more rapid with
monkeys than in this subject, and it is of course possible that it
may have been a sporadic case communicated by human agency.
88 DR. A. E. BROWN ON THE [Jan. 12,
It cannot be said that the results reached with lemurs are
equally exact. Indeed, at the present time, we are not disposed
to rely much upon the charts in this group, except in cases of
positively bad reaction. The greater frequency of doubtful cases
here, than with monkeys, is primarily due to the greater amount.
of normal individual irregularity, especially in regard to the night
drop. ‘This is true even of diurnal species, as the genus Lemur,
in which it 1s sometimes well marked, at others absent. ‘To.
illustrate this two post-injection curves are given in Chart D:
(text-fig. 4, p. 87), of a female Lemur varius, on the first of which,
taken in April 1907, had she been a monkey she would have been
unhesitatingly condemned. In July she gave a good chart,
differing from monkeys only in somewhat advancing the hour of
the night minimum, She died September 8th, without any trace.
of tuberculosis.
There is some reason to believe that on the whole lemurs are:
less susceptible to infection through the usual channels than
monkeys, though by no means immune.
At the present time, after three and a half years of systematic
observation, the findin gs we believe to be justified by the fact that
more than a year has passed without evidence of the existence of
tuberculosis in the Monkey House, are these :—
1. In every case where a positive reaction has followed injec-
tion we have found tuberculosis on autopsy.
2. In negative charts, where no rise of temperature appears,
nor perturbation of the daily rhythm, we have had no reason to.
believe that the disease exists. It has been absent in every such
case after a forty-eight hour observation, where animals have
subsequently come to autopsy.
3. In intermediate charts, neither positive nor negative, safety
lies in nicety of judgment on the part of the observer, this being
mainly a matter of experience. But a very small percentage of
such cases have been tuberculous.
4. In very advanced cases of general tuberculosis the usual re-
action does not always follow injection, but we have found such
cases to be indicated bya notable drop in temperature, and proved
by early death.
The difficulties in the way of applying the temperature reaction
test generally throughout a zoological collection are obvious. The
effort, risk, and various uncertainties involved in the frequent
handling of large and powerful ungulates or carnivores, to
procure the indispensable data, are so great that it is probably
impossible.
Tn this aspect the eye test of Calmette, and skin tests, such as
that of Ligniéres, seemed to be of some promise; but it is un-
fortunately true that experiments made by the present pathologist,
Herbert Fox, M.D., with these tests upon monkeys known to be
tuberculous, have not fulfilled expectations.
We are naturally led to the general subject of mammalian
1909. ] TUBERCULIN TEST IN MONKEYS. 89
temperatures, and a few notes may be given consistently with the
purpose of this paper, together with a list of temperatures that
have been taken with a view to possibly more extended work with
tuberculin.
By artificially causing monkeys to turn night into day in all
that concerned their activity, Simpson and Galbraith reversed the
daily curve of temperature, the low point occurring in the after-
noon, the high point in the early morning. This result, together
with the irregularities we have found in diurnal lemurs, led me to
observe the course in two nocturnal species : Perodicticus potto, in
which the record for twenty-four hours proved to be: 3 P.M.,
OAC (ee vee 99:22- il pew, LOO? 5 tou Aa 90c8 ain ieAeMe,) OO:
11 a.m., 97°6°; 3 p.m., 97°8°: and Galago senegalensis, in which
the 3 p.m. record was 100°6°, the 3 a.m. 102°6°. A single observa-
tion previously made on Galago crassicaudatus at 3 P.M. gave
98°8°. This being low for primates, it is probable that it is near
the minimum point in the daily curve of this species also, and
that the record obtained from the Potto represents the normal in
nocturnal lemurs, corresponding in its main features with Simpson
and Galbraith’s reversed curves in monkeys, the chief difference
being that both high and low points come rather earler.
It seems, therefore, that the temperature drop in primates is an
easily induced physiological result of decreased activity, which
reaches its lowest limit during the period habitually allotted to
sleep, and that the above condition should also be true of the
night-monkeys, Wyctipithecus, but conclusions from five records
taken from two examples of JV. trivirgatus, about 3 P.M., are
obscured by their irregularity. These were 102°2°, 101-5°, 100°,
99-8°, 99:4°. Further investigation of the complete curve in this
genus will be of interest when occasion offers.
The temperatures following were taken with tested thermo-
meters, and nearly all were checked by three observers. Whether
or not, or to what extent, they were affected by excitement, as in
monkeys, can be determined only by a more extensive series of
observations than we have yet been able to undertake. The
records were taken between 10 A.M. and 4 p.m. The temperature
of the building in which the kangaroos were kept was 50°, the
others ranged from 63° to 67°, but, excepting primates, within
these limits neither the hour nor the surrounding temperature had
any perceptible influence in the many cases where the same
animal was tested repeatedly under different conditions.
Bearing in mind, however, the reversed daily curve in night-
lemurs, it is significant to observe that animals of known nocturnal
habit, such as Cercoleptes, Arctictis, Paradoxurus, and Mephitis
among carnivores, and Peromyscus, Dipodomys, Capromys, and
Dipus among rodents, give afternoon temperatures distinctly
lower than is common in diurnal species of their respective
orders.
90 ON 'THE TUBERCULIN TEST IN MONKEYS. [Jan. 12,
| Number of : .
| \Sqesnnens, Maximum. Minimum.
PRIMATES.
| Nyctipithecus trivirgatus ¢ : crodedeae 2 102°2° 99°4°
Hapale sacehius ak wees. : 2 102'8 102
| Perodicticus potto 3 nto dae eae il 97°4:
Galago crassicaudatus @ .................. | il 98°8
a Senecallemsisiidy eee ereeetncneersee: iL 100°6
CARNIVORA.
Nasua rufa ¢ 1 102°6 98°8
» Narica g eae ee 1 100°8
Procyon cancrivorus ay Pog nacee cee coc dee | 2 100°2 99:2
53 lotonruaeeee: Resacaaaiett| il 103°4
Cercoleptes caudivolvulus a Bea ro | 1 975
Bassaris astutus ¢ 1 100°2 99
Viverra megaspila ¢ 1 100°5
Viverricula indica 9 ; il 102
Arctictis binturong g ....2.........4..... 1 99:2
iParadoOxuUnUssnicerlG ue meester ee err ee | 1 98°8
her apie oditus ae stg’ 1 978 |
Nandinia binotata come | i 101
lenpesties mune ond eee seers eee il 103
Mustela furo 2. eed chi na ah ire a 1 102°2
(Galbieing logmlogney, GPP scoconcse case asec 2 1056 | 1042
| | wun, © nba neman ete neeeeE il 101
Mephitis mesomelas Te ? 2 97°8 97°4
Angora Cat ¢ .. ahs etecte at hs iL 1OV2 |
RODENTIA. |
| Dipodomys spectabilis f ............... 1 99°4
# ambiguus J ine, ici ck ae il 96'8
Dipus jaculus ¢ ; LE Er a st 1 99°2
Peromyscus texensis e ASHE Teese stl 1 97°8
Capromys pilorides ¢ .. 1 984 | 95°6
Myopotamus coypus Recah eee 1 TOES) 99
DAS OROCHA, BLAS Gi cavacannasauocoesnseseec | ik 10274 | 102
5 BEymnblaplie 3 a gate | g 102°2 101°6
op agouti g 2. Se aeaee| ) 1OZ:2) )) ) L00;2
33 acouchy ? 1 102°2 1015
PROBOSCIDEA. |
Elephas indicus old @ ..................64. | iL 97°5 96:2
. » oll 2,. Repeated 1 97°5 97-2
a s young Ot BER fice? | iL | 97°8 96°4
| UNGULATA. |
| Wilavananey Tut) Qs jeoocon soo csonaoeco sen ace! 1 | 100'8
Clarins mnGAAG B s.ooc0 ose sco00s sent0n 00a ik | O14
| Anoa depressicornis g ...........--..000 0 | 1 102
_ EDENTATA. |
@holoepus\hofimaniny cesses eee ed 1 94-2
IDS DUS SEXOONCHIS OS sooonsanscnenosenb see | 1 93°4 90
ss novemcinctus ¢ 2 2 93 916
| MARSUPIALIA. |
Didelphys virginiana ¢ ..................| il 94-4, 90°8
Dasyurus maculatus ¢ Q ...........--. 2 93°8 90
Phalangista vulpma @ 9 ............ os) 4. 978 91-6
WER OMS ereemMieMs SL oodocecndessaco-00 il 96°8 944.
2 THOLOUSNUS 6) DL ceccsosoasancancea|| 2 95°2
9 THEMIGONIGISS Z cos cop nocons concen nenl| 1 96°4
MONOTREMATA. |
Hehidmnayacul eatiayg/m sic eeen-eeeee ee 2 87 86
as R Owen (P. Z.S. of London, 1845, p. 81) gives the temperature of the echidna
as 80.
Lowe “Os sjetue q = seq : Ss PW IW IED) VNY IVa
IOEE Tel GOGCI “SZ ‘al
Ae me ae
ie
‘SIIWLIW TS WINS TWEl
Rl mo Ss]stue qd Q aed
IWEK “Tel “GOGT “S “Z “al
pT wossjaureq » oeg “SJL ON US) WIN INL
MAXX Id GOGIT SZ aq
1909. } ON THE WHALE BALAINA GLACIALIS. 91
5. A few Notes on the Whale Balena glacialis and its Capture
in Recent Years in the North Atlantic by Norwegian
Whalers. By Prof. R. Cotter, F.M.Z.S8.
{| Received December 12, 1908. |
(Plates XXV.-XXVII.*, and Text-figure 5.)
Whales captured 18S89-1908.—In our own day, the Nordkaper
is not known to have been observed within the Norwegian area.
Since 1899, however, Norwegians have intermittently carried
on whaling of this species in the sea to the east of Iceland and
round the Faroe Islands, the Shetlands and the Hebrides, where
the Nordkaper has appeared almost every summer in small
companies, or sometimes in large schools,
In the course of twenty years, namely from 188% to 1908, Nor-
wegian whalers have in this way captured about 80 of these
Ww ha ales in the above-mentioned parts of the North Atlantic.
The numbers in the different years have been as follows :—
1889. In April the first specimen was taken on the European
side of the Atlantic, to the east of Iceland. This was a
female, with a total length of 43 feet (13°1 metres); the
length of the head was 3450 mm. (thus being nearly one
fourth of the length of the body).
1890, Seven specimens were taken round Iceland; a skull of
one of these (total length 3920 mm.) and some baleen
being preserved in the Christiania. Museum.
PSO lam all 10 specimens were killed off Iceland, several of them
being caught about 50 miles to the west of that island.
(Skeletons of these are in the Christiania and Bergen
Museums.) One of them was a male, having a_ total
length of about 47 feet (14:4 metres), the skull measuring
3910 mm. in length, or rather more than one fourth of
the entire length. of the body. Another was a female with
a total length “of 43 feet (13:1 metres).
1892. One specimen captured off the Faroe Islands.
1894. Two specimens captured off Iceland.
1897. Two specimens captured off Iceland, one of them Neh
a female with a length of 46 feet (14 metr es).
1898. One specimen taken near the Faroe Islands (a peuitite
accompanied by a young one).
1902. One specimen taken off Iceland.
1903. One specimen taken off the Faroe Islands, and - three
specimens off Iceland, four altogether. Among these
* For explanation of the Plates see page 98.
92 PROF. R. COLLETT ON THE [ Jan. 12,
last was a female, taken in August (length 54 feet, or
16-4 metres), which contained a foetus about 1 metre in
length; another was scarcely more than a half-grown
young one.
1904. Two specimens were captured to the south-east of Iceland.
1905. (One was wounded off St. Kilda, but escaped.)
1906. Six killed, and more seen (in company with Balenoptera
borealis) off the Hebrides, between the 13th June and
the 4th August.
1907. A large number seen, and 24 killed off the Hebrides.
The latter were of both sexes; all the females were
gravid.
Two specimens were also taken off the Faroe Islands,
making 26 in all.
1908. Several hundred seen, and 20 killed off the Hebrides,
between June 18th and July 9th. Those killed were of
both sexes; none of the females were gravid.
Five specimens were also captured off Inishkea, Ireland,
between June 8th and June 13th (among them one female
and a young one), making 25 specimens in all.
Whaling in the Hebrides, 1906—1908.—The largest capture of
Nordkapers in the present day took place during the past three
years, when a single company (at Station Buneveneader, Harris),
came across large schools of them in the sea off the Hebrides,
and brought back 50 (6 in 1906, 24 in 1907, and 20 in 1908).
About these I have received some particulars from the manager,
Capt. Carl Herlofsen.
The six specimens in 1906 were taken on June 13th and 15th,
July 18th and 31st, and August 4th (two on July 31st).
In 1907, the first two specimens were captured on June 13th.
On June 15th another two were taken, and the rest later in the
same month, ten in all; in July fourteen were captured, the
last on the 26th July. The 11th July was a successful day, six
whales being captured, four of them in the course of six hours.
In 1908, on the 13th and 22nd June, two specimens were
taken, the remaining 18 being taken between the 3rd and the
9th July. On two occasions five whales were killed in one
day, namely on the 4th July (all males) and on the 7th July
(three males and two females).
The five specimens that were killed the same year off Ireland
had probably been on the whaling-ground at the beginning of
June. On the first day of whaling, the 8th June, one Nordkaper
was taken, and the remaining four were caught within the
next few days. They were all separate, and no schools were
observed.
Size-—Among the 24 specimens captured in 1907, males and
females were in equal numbers; while of the 20 in 1908, twelve
were males and eight females.
1909. | WHALE BALANA GLACIALIS. 93
The specimens of the latter year averaged rather smaller
than those of 1907. The females seem to be larger on the whole
than the males; and the largest of all the specimens was a
female, whose length was 50 feet (15-2 metres).
Text-fig. 5.
Balena glacialis, male; from the Hebrides, July 6th, 1908.
The 24 specimens in 1907 were of the following length :-—
Mauss (12). FEMALES (12).
English feet. English feet.
48 (14°6 metres 49 (14°9 metres)
475 (144 ,, | ANS (IPA,
Aan (as dere (fee
AGES WGC ss ASN (GMa
Agee (GA ow A7b (aan |,
46 (14 33 47 (1433,
ABR O (RRS 1 5 A sates
ASR GIB Os. Le tale eas
45 (3s 46 (14 »
AS) Bes AL SS ls
Tey (ee | A ase
)
)
)
AG (ESS, | 475 (144,
)
)
)
)
)
aS wes OS OS ESS
94 PROF, R. COLLETT ON THE [Jan. 12,
Most of the 12 males thus had a length of from 46 to 47
feet, the average being 45:8 feet (13°9 metres).
Among the 12 females, the length was most frequently between
47 and 48 feet, the average being 47 feet (14:3 metres). In
girth, the males measured 33°7 feet, the females 36°6 feet *.
The 20 specimens in 1908 were of the following lengths :—
Mates (12). FEMALES (8).
English feet. English feet.
(14:3 metres 50 (15°2 metres
AT (14:3 Cn, 49 (Eg
(
=
oo
—
=
is)
pm
»
S
SoS SS Sr
)
)
)
AG (yg Bye GS
)
)
)
)
)
Thus in 1908 the length of the 12 males was between 36 and
47 feet, the greater number of them being from 42 to 46 feet.
The average length was 43:8 feet (13°3 metres).
Of the 8 females, one was 50 feet in length, and another, a
young one, 31 feet, the remainder being between 43 and 49 feet.
The average length this year was only 44 feet (13°4 metres).
The females in 1908 measured on an average 35 feet (10°6
metres) in girth, and the males 33°8 feet (10°3 metres), but the
girth varied considerably. Among the males this year, there
was one specimen that had a total length of 46 feet, and
measured exactly the same in girth, namely 46 feet. Although
it was not greatly inflated with gas (it had been dead only
twenty-four hours), it was almost as round “as a ball” when it
lay on the beach.
Colour.—There is no great difference to be found in colour
between male and female.
A uniform black must be considered to be the typical colour,
covering the entire body without any great differences of shade.
In some specimens, however, more or less of the ventral surface
was white. The boundary of this white colour was clearly
defined against the black sides; in many specimens the white
area was somewhat constricted in the middle, and in parts,
especially towards the sides, was covered with oblong, black
spots.
* Haldane, Ann. Scot. Nat. Hist., April 1908, p. 69.
1909. WHALE BALAINA GLACTALIS, 95
The white-coloured belly occurred in both males and females.
Of the 6 specimens captured in 1906, one adult specimen and
one young one were white-bellied. Out of the 24 captured in
1907, six specimens, or one fourth of the whole number, were
white-bellied. Of the 20 specimens captured in 1908, only two
were white-bellied (one male and one female). Thus 20 per
cent. of the 50 specimens captured in the course of the last
three year's have been white-bellied.
None of the five specimens caught in 1908 off Iveland was
white-bellied. The specimens were of both sexes, and the largest
measured about 50 feet in length.
In the black colour on the belly in most, though not all, of
the specimens, a large number of white stripes occurred, running
in all directions, and measuring up to one metre in length and
about 50 mm. wide. It is possible that these stripes may have
been produced by the rubbing of the animal against the bottom
when following the plankton-crustaceans upon which it feeds,
and that they are of the same origin as those described in certain
old specimens of Mesoplodon bidens and others.
The pectorals were black on the whole, both in the black- and
in the white-bellied specimens, though often with a faint white
marbling on the upper surface and the margin.
The baleen is black, both in the white-bellied specimens and
in the black, though in some specimens a few of the foremost
plates were white.
The number of plates was stated to be about 250 on each
side. The bristles were also all black, and almost as fine as
silk. One of the longest plates, which was presented to the
Christiania Museum, measured 2225 mm. in length; the bristles
were longest at the tip, where their length was 450 mm., but
elsewhere measured about 250 mm. The longest plate that has
been measured (Iceland 1903) had a length of 2700 mm.
The peculiar wart-like excrescences are situated in irregular
rows along the upper and lower mandibles. The largest of
these, as in B. australis*, are at the tip of the upper jaw, where
several sometimes join, and together form the largest ‘bonnet ”-
then one on each side of the tip of the lower mandible, and
finally one immediately above each eye.
The value of a Nordkaper at the present time is from about
6000 to 10,000 kroner (£330 to £550). The blubber, which
in some specimens is of a pale pink colour, has a thickness of
about 260 mm. The amount of oil it contains varies from 10 to
30 barrels (of the first quality).
In the two young specimens mentioned above as captured in
1907, the blubber was pure white; the animals were exceedingly
fat, and yielded about 30 barrels of oil each.
The weight of baleen in a full-grown specimen is from 250
* Lénnberg, Kgl. Sv. Vetensk.-Akad. Handl. B. 40, no. 5, p. 45 (Oct. 11, 1906).
‘96 PROF. R. COLLETT ON THE [Jan. 12,
to 330 kilogrammes, and is valued at abgut 6800 kroner (£375).
From four full-grown whales, about one ton of baleen is obtained,
which will thus fetch about 27,000 kroner, or £1500*. A
single one of the longest plates of whalebone 1s worth about
38 kroner (£2 2s.).
Habits —Every year of the whale-fisheries in the Hebrides,
the whales kept almost to one place, always occupied in seeking
food among the pelagic crustaceans. In 1906 they stayed
nearly seven weeks, but they were then more scattered, and
appeared more irregularly.
In 1907 they were on the ground for about six weeks, during
which time they appeared sometimes separately, sometimes in
small schools. The school that took up its quarters in this spot
in 1907, consisted of at least 100 whales.
In 1908, the plankton-bearing currents probably flowed nearer
land than in 1907, for the whales might be met with quite in
the shallow water between islands and rocks. Their stay this
year was of only three weeks’ duration.
The schools this year consisted of several hundred, and, as
-already mentioned, the boats of the Station several times captured
from two to five whales a day.
In 1907 the school was unaccompanied by any other species
of whale; but in 1908 they came with hundreds of Rudolphi’s
Rorquals (Balenoptera borealis), which were just on their way
morth 7.
The five specimens killed off Ireland in 1908 were also
accompanied by B. borealis.
The Nordkaper is not timid, and is on the whole easy to
approach. The harpoon used is a bomb-harpoon of the kind
used in the Arctic Ocean. As the blubber is of considerable
thickness, the harpoon should if possible be discharged at close
quarters. If it strikes in the right place, the whale soon dies ;
but if it is only wounded, it becomes very violent in its
movements, to the no small danger of the boats, although it
-does not attack them; it plunges round in the water like a
ball, and often gets the line wound several times round its body.
Notwithstanding the thick build of its body, it is able to bend
it until the head nearly meets the flukes.
Tt is fond of lying quietly on the surface of the water; and
it moves slowly, with its blow-holes above water. The jet these
sent up could be seen from a considerable distance, and was
about 5 metres in height. It was comparatively thicker than
that of a Common Rorqual (B. physalus); a closer view shows
it to be distinctly formed of two jets falling to different sides,
* The 24 specimens caught in 1907 yielded a total of rather more than 6 tons
of baleen, of which the value was more than £9000 (163,000 kroner).
+ B. borealis (Norwegian “Sei-hval”) appears annually in larger or smaller
numbers in the plankton-currents off the coasts of Tromsé and Finmark, generally
staying from the middle of June to the middle of August.
1909. } WHALE BALENA GLACIALIS. 97
As a rule, it blows five or six times in succession, and then
remains under water for from ten to twenty minutes.
It dives almost perpendicularly, and therefore in diving shows
the whole of the flukes. It sometimes leaps high in the water,
but it has never been seen to leap quite out of the water.
It has never been heard to make any sound.
Its food, both in the Hebrides and off Iceland, was found to
be exclusively pelagic crustaceans (the “krill” of Norwegian
whalers), a Euphausiid about half an inch long, probably
Boreophausia imermis.
Parasites.—All specimens were infested with thousands of
Cyamus (the “lice” of whalers), which are especially found in
the furrows of the excrescences along the jaws. They may also
occur around the genitalia and scattered over the body.
Young.—Among the numbers that have frequented the waters
round the Hebrides during the last three years, no small young
ones were found. The three smallest captured had total lengths
of 31, 36, and 37 feet (9:45, 10-9, and 11:2 metres). One of
these young ones was white-bellied, and its baleen was of a
lighter shade of colour than that of the black-bellied.
Propagation.—Three specimens were observed just before
copulation on the 7th July, 1908. A female was lying on her
back, and on each side of her lay a male with extended genital
member, when the vessel came upon them and secured the female.
The twelve females killed in 1907, in June and July, in the
Hebrides, were all gravid. The feetuses were all more or less of
the same size, having a length of from 1 to 14 metre; their
colour was always pale blue, with no trace of white on the
under surface.
In the largest foetus, the first rudiments of baleen had begun
to appear.
In 1908, eight females were killed in the Hebrides, but, as
previously stated, none of them was gravid. It is therefore
possible that the gravid females go in separate schools,
There is also an account of a foetus of about 1 metre in length
found in the summer of 1903 off Iceland (in a female 54 feet in
length, or about 16-4 metres).
From the above observations the following facts appear :—
Balena glacialis may at present be met with in the summer
in the North Atlantic in schools of 100 or more.
The length of most of the full-grown males captured in the
years 1906-1908 was from 46 to 47 English feet (up to 48 feet), or
from 14 to 14°3 metres (up to 14°6); that of the females generally
from 47 to 48 English feet (up to 50 feet), or from 14:3 to 14°6
metres (up to 15:2 metres).
The greatest length was that of a gravid female, and amounted
to 54 feet, or 16-4 metres (Iceland, 1903).
Of the 50 specimens captured in the summer of 1906-1908 in
the Hebrides, about 10 per cent. were white-bellied.
Proc. Zoou. Soc.—1909, No. VII. ie
98 MR. R. E. HOLDING ON THE HORNS [ Feb. 2,
Copulation may take place in the beginning of July, and
gravid females have been found in Juneand July, with foetuses of
about 1 metre’s length.
Their food in the summer months in the North Atlantic is.
(as far as has been observed) plankton-crustacea.
EXPLANATION OF THE PLATES.
PratE XXYV.
Balena glacialis, male, from the Hebrides, July 8, 1908.
PratE XXXVI.
Balena glacialis, male, from the Hebrides, July 6, 1908.
Pratt XXVII.
Balena glacialis, female, white-bellied, from the Hebrides, July 6, 1908.
February 2, 1909.
Freperick Ginter, Esq., Vice-President, in the Chair.
Mr. C. Tate Regan, M.A., F.2Z.8., exhibited specimens of the
Char of Lough Melvin (Salvelinus grayi, Gimth.) and of the
Char from a little loch under Ben Hope, Sutherlandshire, recently
described by him under the name Salvelinus maxillaris. He
pointed out the differences between the two forms, and called
attention to the interest attaching to the study of this too much
neglected group of British freshwater fishes.
My. R. E. Holding exhibited several skulls and photographs of
the St. Kilda or Hebridean Four-horned Sheep, and made the
following remarks concerning the horns in this variety :—
“That there are several well-defined breeds of the domesti-
cated Sheep which carry normally four horns is now well known.
Explanation of Text-fig. 6 (see opposite).
A. Head of St. Kilda Ram, showing wide type of the horns. Length of upper horns
182 inches each.
B. Ditto, showing the narrow or more contracted type. The upper horns forming
almost a complete circle; in both specimens the lower horns had to be cut to
prevent their growing into the jaw.
C. Photograph from life (by E. M. Machugh) of an exceptionally good head of a
well-known Scotch black-faced four-horned breed, which was established some
thirty years ago from an odd Ram Lamb which occurred in the flock. The
‘ owner taking some interest in the matter, was enabled by selection to cause
the variety to become permanent, some remarkable heads being the result.
D. Skull from the same flock as B, showing two separate horn-pedicles or supports
(1), covered by one sheath (2), indicating that these supernumerary horns
have their origin in duplicated centres of the frontal bone. The upper left
poimting backwards is an unusual variation.
E. Upper part of the skull of a St. Kilda Ram Lamb at six weeks old, showing an.
early stage in division of the bony pedicle which supports each horn. QO, occi-.
pital; P, parietal; F, frontal; N, nasal.
1909. | OF THE ST. KILDA FOUR-HORNED SHEEP. 99
Text-fig. 6.
Heads etc. of Four-horned Sheep.
100 ON A FIGHT BETWEEN A WHALE AND A SworDrFIsH. [ Feb. 2,
The St. Kilda or Hebridean and the South-African varieties
are the better known, but there are also an Indian variety, of
which several specimens are living in the Society’s Gardens,
having some slight variations, and a local breed of the well-known
Highland ‘“ Black face,” which also bear four horns. The
St. Kilda is entirely black, the fleece being dark brown; the
South-African form is piebald on the face and legs, with black
irregular marks on the fleece; the Indian has spotted face and
legs and white or grey fleece; and the Scotch variety has charac-
teristic face-markings and long-stapled wool.
“Although at first there seems to be a somewhat perplexing
irregularity i in the form and pitch, as it were, of the horns of
these Sheep, there can, I think, be seen a fairly constant type
which separates the horns of the St. Kilda Sheep from those of
other varieties. Upon looking over a considerable series, it is
apparent that there are two distinct types—one in which the
median horns are directed well forward ina semicircular curve, as
in A, text-figure 6 (p. 99), and the other, B, in which the median
horns take a much smaller curve over the face. I have not noticed
an intermediate stage in this breed. The lateral horns spring at
almost right angles from the skull, and grow at times so close
inwards that the points would penetrate the skull if notcut. The
median horns of the South-African variety are usually erect,
or, if curved, take a backward inclination, rarely directly ‘forwar d
as in the St. Kilda form. ‘The lateral horns are also of wider
curve, and so grow clear of the head. The Highland variety
follows very closely the Hebridean form, as shown in C.
“Tn the same flock of St. Kilda Sheep some variety may occur
in the number of the horns, as indicated by D, which come from
the same source as A.. This is apparently due, not to any arrest
in the growth of the bony horn-bearers, but to their fusion under
a single horn-sheath, as indicated by the specimen. These super
numerary horns are not due to an antero-posterior cleavage, 7. €
from apex to base, but to segmentation of the centres of ossifica-
tion of the frontal bone, as indicated by the skull of a Lamb, H,
at six weeks old.”
Mr. Malcolm Maclaren, through Mr. C. Davies Sherborn,
F.ZS., called the attention of the Fellows to an account of a fight
between a Whale and a Swordfish observed by the crew of the
fishing-boat ‘ Daisy’ in the Hauraki Gulf, between Ponui Island
and Coromandel, as reported in the ‘ Auckland Weekly News,’
19th Nov., 1908. A cow whale and her calf were attacked by a
12 ft. 6 in. swordfish, the object of the fish being the calf. The
whale plunged about and struck in all directions with her flukes.
Occasionally the fins of the swordfish were seen as he rose from
a dive, his object apparently being to strike from below. For
over a quarter of an hour the whale circled round her calf,
lashing furiously and churning up the water so that the assailant
was unable to secure a good opportunity for a thrust. At last,
1909. ] ON THE FAUNA OF CHRISTMAS ISLAND. 101
after a fruitless dive, the swordfish came close up and made a
thrust at the calf, but received a blow from the whale’s flukes
across the back, which apparently paralysed it. It was killed
and hauled on board the boat without difficulty, while the whale
and calf went off towards Coromandel with splashings and
plungings. The whale’s blow had almost knocked off the back
fin of the swordfish and heavily bruised the flesh around it. No
threshers accompanied the swordfish.
Dr. C. W. Andrews, F.R.S., F.Z.S., gave the following account
of his visit to Christmas Island in 1908, and illustrated his
remarks with lantern-slides :—
“Christmas Island in the Indian Ocean (S. lat. 10° 25’, E. long.
105° 42’) lies some two hundred miles south of Java, which is the
nearest land. It has an area of about forty square miles, and its
highest point is 1200 feet above the sea-level. The island con-
sists mainly of coral-limestones resting on a basis of volcanic rock
interstratified with foraminiferal limestones; the whole surface
is thickly covered with forest and jungle. The fauna and flora
of this isolated spot have been the subjects of numerous papers,
a list of which up to the year 1900 is given in the ‘ Monograph
of Christmas Island,’ a volume which contains a detailed account
of the collections Gritieh were made by me during my former
visit in the years 1897-8. Since that date the island has been
visited by Messrs. Ridley and Hanitsch of Singapore, and the
former has published an interesting account * of the flora, to
which he was able to make a considerable number of additions.
Almost up to the date of my first visit the island had been
uninhabited, and was only visited by ships on very rare occasions ;
but since then, owing to the quarrying and export of the valuable
deposits of phosphate of lime, a considerable population has been
imported, and many ships call either to bring stores or to ship
cargoes. The consequence of this is that many animals and
plants have been introduced from time to time, and it was for the
purpose of investigating the effect of these introductions on the
native flora and fauna that, at the suggestion of Sir John Murray,
T revisited the island in the autumn of last year, remaining there
about three months. The changes that have taken place are, as
might be expected, chiefly noticeable in the immediate neigh-
bourhood of the settlement and quarries, while the rest of the
island, although traversed by roads in several directions, is
practically unchanged. To this general statement there is,
however, at least one important exception, and that is that the
two species of native rats seem to have become totally extinct.
At the time of my former visit these animals swarmed over the
whole island, one, Jus macleari, being found practically every-
* “The Botany of Christmas Island,” Straits Branch, Royal Asiatic Society
Journal, June 1906, p. 121.
102 ON THE FAUNA OF CHRISTMAS ISLAND. [ Feb. 2,
where; the other, J/us nativitatis, more adapted for burrowing,
was for the most part confined to the higher ground. Last year,
in spite of continual search, not a single specimen of either species
could be found in any part of the island. This complete disap-
pearance of two such common animals seems to have taken place
within the last five or six years, and to have been the result of
some epidemic disease, possibly caused by a trypanosome, intro-
duced by the ship-rats. These are a variety of Mus rattus, and
have been introduced in considerable numbers, though they do
not seem to have spread to the remoter parts of the island at
present, at least to any great extent. The disappearance
therefore of the native forms cannot be due to direct competition
with the intruders, but must be the result of disease, a con-
clusion supported by an observation made by the medical officer,
Dr. McDougal, who told me that some five or six years ago he
frequently saw individuals of the native species of rats crawling
about the paths in the daytime, apparently in a dying condition.
Thope that it will be possible to obtain information as to whether
any trypanosome or other pathogenic organism occurs in the blood
of the imported rats.
‘Of the other native mammals the shrew (a variety of Croct-
dura fuliginosa) is probably also extinct, at least no specimen
was either seen or heard during my visit. The large fruit-bat
(Pteropus natalis) is more numerous than formerly, at least near
the settlement, probably in consequence of the much larger
number of fruit-trees now to be found there. Cats have been
introduced, and are becoming numerous; in some cases they
have taken to the woods, and occasionally cause great destruction
among the poultry which are reared in large numbers.
‘The native land-birds all seem to be as numerous as formerly.
The large fruit-pigeon (Carpophaga whartoni), which is. used
for food, is protected during the breeding-season, and at other
times the number killed is limited so far as possible. These
birds have not yet acquired any fear of man, and allow themselves
to be snared by hand as easily as ever. The frigate-birds do not
nest in the neighbourhood of Flying Fish Cove in anything like
such numbers as formerly, but are still very numerous elsewhere.
The yellow tropic-bird (Phaethon fulvus) seems to have increased
in numbers. No introduced bird has obtained any footing on
the island.
“No changes of importance were noticed among the native
Invertebrates; but the large Scolopendra, of which a very few
individuals were noticed during my first visit, is now much
more numerous. Probably many insects have been introduced,
but until my collections have been fully worked out, nothing can
be said on this point.
““ A large number of plants have been introduced, and probably
the clearing of the forest and cutting of roads will enable them
to spread much more rapidly in the future than they have done
in the past. Already the papaia and chillies have extended
1% FAs So ISOS), Jil, 2OMvalals
PHYLLIUM CRURIFOLIUM. POON ee Ace
1909. } ON THE LIFE-HISTORY OF THE LEAF-INSECT. 103
their range considerably. In the clearings in Flying Fish Cove
and on the shore-terrace it is interesting to note that nearly all
the secondary growth consists of a small tree, Jelia azederach,
which at the time of my first visit was represented by two or
three examples only, and these had most likely been introduced
by man. In other parts of the island clearings are often
occupied by great numbers of seedlings of Jnocarpus edulis ;
this is one of the consequences of the extinction of the rats,
which formerly fed largely on the fallen fruits ef this tree.
“Much time was spent in collecting the marine fauna, espe-
cially of the reef at Flying Fish Cove, and I hope that reports on
some of the groups will be published shortly.”
The following papers were read :—
1. Preliminary Account of the Life-history of the Leaf-
Insect, Phyllium crurifolium Serville. By H. 8. Leran,
Honorary Research Fellow in the University of
Manchester.*
[Received December 23, 1908. |
(Plate XXVIII. 7)
The present paper is intended to form a preliminary account of
the life-history of a Leaf-Insect, Phyllium crurifolium? Serv. from
the Seychelles Islands. I have had this species under observation
during the past year, and although I have been able to note
several facts of interest in connection with the development,
many of the more important details remain to be worked out.
It was thought, however, that notwithstanding the fragmentary
nature of my observations up to the present time, some account
of this remarkable insect might prove interesting, and it is on
this consideration that the following preliminary account has been
written.
My study of P. crurifoliwm has been carried on in the Zoological
Laboratories of the Manchester University, and also in the hot-
houses at my own residence in Worsley.
I had neither sufficient material nor apparatus for making a
complete study of the insect and its development during the past
year. The work will be continued, however, during the coming
season, and with abundant material at my disposal I hope to com-
plete the details of the study and publish the results in a fully
illustrated memoir.
During the autumn of 1907 I received a hundred ova and some
* Communicated by Professor S. J. Htcxson, ¥.B.S., F.Z.S.
+ For explanation of the Plate see p. 113.
+ Kirby (1904) ineludes this species in his genus Pulchriphyllium.
Que MR. H. S. LEIGH ON THE | Feb. 2,
thirty young larvee of Phylliwm crurifolium from Mr. St. Quintin,
F.Z.S., and it is owing entirely to his kindness in presenting them
to me and also in furnishing me with a few of the more important
suggestions as to the conditions necessary for successfully rearing
them, that I have been able to observe the habits of the insects in
confinement.
The eggs that I received were laid by insects which Mr. St
Quintin had reared during the winter of 1906 and 1907, and he
is, I think, the first naturalist who has bred the Leaf-Insect im
this country. The original stock was brought from the Seychelles.
to England in the spring of 1906 by Lord Crawford.
The genus Phylliwm*, which constitutes the tribe Phylliides and
raeladee all the Leaf- Tensace, comprises perhaps some of the most
extraordinary of all living insects; their wonderful similarity to.
vegetable structures has often aroused the admiration of natura-
lists, but it is only upon gaining a more complete knowledge of
their life-histories that we can fully appreciate this most as-
tonishing example of ‘“ Protective Resemblance.” It is not a
matter of much surprise that the Leaf-Insects were thought at
one time to be of a vegetable as well as of an animal nature.
This imaginative belief has given rise to several peculiar ideas,.
and one prevalent notion amongst the people in those countries
which these insects inhabit is, that the insect is in reality a
changed leaf that has adopted the strange practice of walking (de
Borre, 1883).
The genus of Leaf-Insects was thought in the time of Linnzeus
to consist of only one species; this was named Mantis siccifolius:
(1767), and was figured by Rdésel (1749). Stoll (1815) gave
figures of two species, whilst Serville (1839) mentions three species
in his ‘ Histoire Naturelle des Orthopteres. In 1843 Gray
described thirteen species, many of which were new, and West-
wood (1859) mentions fifteen species. Joly (1871) described more
fully than any previous author the natural history and anatomy
of Phyllium crurifolium Serv.; and later, in 1887, Brongniart
gives a short account of the development of Phyllium siccifolium
Linn. Since that time very little appears to have been written
until 1903, when Morton’s paper “ Notes sur l’élevage des.
Phyllies” was published; and quite recently two short papers
have been written by St. Quintin: one “ Leaf-Insects in
Captivity ” (1907), and the other ‘“‘ Notes on the Life-history of
the Leaf-Insect ” (1908).
Apart from these, little work appears to have been done
recently, and our knowledge of the life-history of the Leaf-Insect
is still meagre.
The first living examples of Phylliwum were brought to this
country in 1854. The bringing of living specimens was attended
with some difficulty, but Mrs. Blackwood, who was greatly
* Kirby (1904) includes three genera in his subfamily Phylliine, viz. Chitoniscus,
Pulchriphyllium, Phylliun.
1909. ] LIFE-HISTORY OF THE LEAF-INSECT. 105
interested in these strange insects whilst in India, attempted to
bring some back with her; she received some eggs in 1854 which
hatched in due course, and Murray (1856) gave an account of a
specimen which was reared to maturity in the Royal Botanic
Gardens of Edinburgh. Prior to this, I believe the trans-
formations and habits had not been watched by any naturalist.
The Phylliums are peculiar to the Old World, being mostly
confined to the islands of the Indian Ocean, and they would seem
to have a special predilection for insular life. Several species
inhabit the Seychelles and Ceyion, whilst others occur in
Mauritius, Borneo, Java, Celebes, and the Philippines, and some
are reported to extend as far as the Fiji Islands. One species,
Phyllium scythe, is recorded from the mountainous regions of
Noxthern India.
The eggs of the Leaf-Insect have been studied in Europe on
several occasions: by Murray in 1855, Joly in 1871, Henneguy
in 1890, and others—they all speak of their great resemblance to
seeds. The egg of Phylliwm crurifolium is about the size of a
sweet-pea seed and resembles very closely the seeds of certain
umbelliferous plants. Murray, speaking of P. scythe, says: ‘If
the edges of the seed of the Mirabilis jalapa were rubbed off, the
seed might be mistaken for the ege.” The egg—which is
in reality a capsule containing the egg—is of a brown colour,
somewhat barrel- shaped, with “five longitudinal ribs, all of w hich
are equal distances apart, except two between which the space 1s
wider. The surface of the egg is rough and cork-like, and there
is usually an irregular row of small pits in the spaces between the
longitudinal ribs. The large space is much smoother and flatter
than the other portion of the egg, and in the centre there is a
groove of an oval form extending almost from the apex to the
base and enclosing a small scar or hollow which I believe has been
compared to the hilum in seeds. At the apex of the egg a
conical lid or stopper is attached, whilst the base is slightly
concave.
The capsule of the egg appears, without the aid of a lens, to be
of a rather fibrous nature, but when examined with a microscope
the true porous texture is revealed. If, as Murray says, this
outer covering had been of a firm subs stance, the embryo insect
could not have received the amount of air and moisture necessary
for its existence. Moisture and warmth are two extremely im-
portant factors in the development of Phylliwm, and certainly if
the eggs were surrounded with a compact substance the young
insect could not develop. As in most Phasmide, the eggs are not
glued on to surrounding objects, but are deposited loosely and fall
through the foliage to the ground where they remain for some
time before hatching; they are retained for a few minutes
between the gonapophyses of the female after their extrusion
from the oviduct and are afterwards shot out to some distance.
In some cases ova were found three feet or more from the
female.
106 MR. H. 8. LEIGH ON THE [ Feb. 2,
This habit of discharging the eggs no doubt ensures a better
distribution of the offspring, as the females are extremely sluggish
creatures, and if the eggs were simply dropped they would in all
probability be clustered together in masses and fall an easy prey
to their enemies.
It is necessary when rearing the Phylliums in captivity to
provide abundant moisture as well as heat for the ova. They
were placed in an orchid house where a temperature varying from
65°—-85° F. was maintained, the atmosphere being also exceedingly
moist.
Under these conditions the ova began to hatch about the
end of August 1907, the first larva appearing on August 28th,
and they continued to hatch very irregularly until the end of
January 1908. Although the majority of the larve appeared
during September, October, and November, I found that their
emergence depended to a great extent upon a high temperature ;
none hatched unless the temperature exceeded 70° F. Morton
and St. Quintin each speak of this irregularity in the hatching of
the eggs, and the latter says that although his larvee continued to
emerge during several months, “ the: ova were all deposited
between the 7th April and the 15th May.”
I think it is very probable that the time passed in the egg
stage 1s exceedingly variable and may be prolonged. In confine-
ment embryonic development requires four to seven months for
its completion. Apparently the larve emerge very irregularly in
the Seychelles, for some were found in the early larval stages side
by side with the adult insects.
The larva, when ready to emerge, pushes off the lid or stopper
to which reference has already been made. When newly-hatched
it is about 16 mm. in length and possesses the characteristic form
of the Phylliums. The head and prothorax are reddish-brown ;
meso- and metathorax brown with a red dorsal line; flattened
expansions of the abdomen reddish-brown, but marked with dark
brown semicircular rings from the fifth to ‘the tenth segments.
I found that the young larva takes no food until four days after
hatching, and during this time it is rather active, moving about
with a singular hesitating and staggering action, which is even
accentuated if the insect is alarmed.
The plant upon which these insects principally feed is the
Psidium guava, but in Ceylon they are said to feed also on the
leaves of the tea and certain species of lemon trees.
This question of food probably offers one explanation for the
young larve fasting a few days, as the chances are it would be
extremely difficult for them (on emerging from the eggs lying on
the ground) to climb up the stems of their food-plant through the
thick and entangled vegetation and obtain food at once.
Not being able to procure any of the natural food-plants, I
followed the example of Mr. St. Quintin and placed my young
larvee on oak leaves which I ional they took to well; they
require, however, to be provided with plenty of moisture, and to
1909. ] LIFE-HISTORY OF THE LEAF=INSECT. 107
ensure this, | sprayed them once a day with tepid water. It was
most interesting to see the thirsty larve eagerly drinking the
drops as they hung from the leaves and branches after the
Spraying.
As the autumn advanced the leaves of the oak became less
nutritious, and it was necessary to find another substitute for the
larve. Again following the advice of Mr. St. Quintin, I placed
the larve on small binges of the evergreen oak (Quercus ilex) and
on this they settled down very well.
The rate of development, 7. e. the time occupied between the
«leposition of the eggs and the appearance of the imago, depends
very largely upon the temperature. TI have not yet been able to
work out the exact length of the various larval stadia, but have
‘determined their approximate lengths, and I should think that
under favourable circumstances the whole development would be
completed in ten or eleven months.
The Leaf-Insect is Paurometabolous and corresponds to the
general rule in the Orthoptera. The larve in all stages are on
the whole similar to the adults and there is no very abrupt
transition, development taking place by a gradual increase in size.
In the newly- hatched larva neither joni nor wings are present,
these being acquired during the later stages, but only attaining
their full dimensions in the imago state. There is no pupal stage,
the larva merely passing through a slight resting stage prior to its
reaching the perfect condition. JI found that there are six or
seven larval stadia, so that the number of ecdyses is six or
seven. Murray gave three as the number of ecdyses in Phyllium
scythe, and if this is correct it appears curious there should be
such a discrepancy in the number of moults in the Phylliums.
The larve moult for the first time when about six weeks
old if kept in a mean temperature of 66° F., the succeeding
moults following at intervals varying from five to six weeks.
The colour of the newly-hatched larva is, as previously stated,
reddish-brown streaked with dull red, but this soon changes and
the larva passes through a regular series of colorations from brown
to green during the first fortnight. In the first day or two of
lereal life hardly any change is mene amie excepting the slightly
paler colour ; but when the insect is a week old a yellowish-green
colour is assumed which gives place in about a fortnight. to a
beautiful pale green on the dorsal and to a rich glossy green on
the ventral side. Later, as the larve increase in size, their colour
becomes more varied and, although green predominates, many
shades occur. Some individuals are beautifully marked with
different shades of brown and yellow. These colours combined
with the positions assumed by the larve render them most in-
conspicuous on the leaves.
Nor is the resemblance to plant-life confined to colour, for the
larva has a very peculiar gait which imitates to a remarkable
degree the shaking of the leaves rather than the movements of an
insect. The larva of the second stadium is about 20 mm. in
108 MR, H. S. LEIGH ON THE [ Feb. 2,
length, but apart from size differs little from that of the previous
stage. The sexes, so far as I could see, were indistinguishable at
this stage.
In the third stadium the external sexual characteristics first
make their appearance. The females are somewhat larger than
the males, the former being 28 mm. in length, the latter about
26 mm., whilst the flattened expansions of the abdomen are
slightly broader in the females than in the males. The future
wings of the male, although at present very insignificant, may
now be seen in the form of two small processes on the metathorax ;
these being entirely absent in the females. Further, the legs of
the male larve are beautifully mottled with irregular patches of
brown and yellow, those of the female being generally without
these decorative markings. The fourth larval stadium is charac-
terised by increased sexual dimorphism ; the male is about 33 mm.
in length and 19 mm. in greatest breadth ; the antenne have now
increased a little in length, being 3 mm.; and the future tegmina
and wings are present in the form of processes about 4 mm. and
2 mm. long respectively. The female is larger and of a somewhat
rounder shape; length 35 mm., breadth 19 mm. The femora of
the prothoracic legs are of wider proportions than those of the
male, whilst the femora of the mesothoracic legs have, in the
middle of their posterior lobe, a brownish spot which is seldom
present in so decided a form in the male larve. The future
tegmina may be noticed in the shape of two small processes,
about 14 mm. in length; the antenne remain short, in contrast
to those of the male. The sexual dimorphism becomes gradually
more pronounced at each of the succeeding stadia, and reaches its
fullest significance in the adult condition. In the penultimate
stage the sexes are very dissimilar; the male*is about 55 mm.
in length, 25 mm. in greatest breadth; has moderately long
antennz ; the future tegmina and wings are well defined as two
pairs of dorsal appendages, and the expansions of the prothoracic
legs are comparatively small. The female is altogether much
larger, being about 70 mm. long and 35 mm. broad; has short
antenne ; only one pair of dorsal appendages (tegmina), and the
foliaceous expansions of the prothoracic legs are very large.
The larve throughout their lives are diurnal, but seem to.
show a decided partiality to feeding during the first and last
hours of daylight; food is taken at other times of the day, but
a decided preference is shown (at all events in the earlier larval
stages) for the twilight. As the sun sinks below the horizon
and the light fails the larve begin operations by a number of
preliminary swinging moyements, after which they move in a
very hesitating fashion in search of food; having discovered
the leaves that are palatable to them, they feed with great.
avidity for fifteen or twenty minutes. At the end of this time
darkness has usually set in and the larve once more settle down
to await the morning, when the same operations are repeated.
I found that the last larval stadium is frequently prolonged
1909. ] LIFE-HISTORY OF THE LEAF-INSECT. 109
and lasts eight weeks in many instances. The last week of this
stadium is spent in a rather torpid condition during which the
larva takes little or no food. A few days before each ecdysis
the larva (as in many species of insects) ceases to feed, and
having secured a good foothold on the underside of a leaf remains
motionless.
After each ecdysis the larva usually devours its cast skin
except the legs, and then fasts for about twenty-four hours before
recommencing to feed upon the leaves. It takes some time for
a moult to be completed, and forty-five or fifty minutes generally
elapse from the time of the old skin splitting to the time of
the larva being entirely free. The whole process of moulting
appears, however, in the Phyllums to be most curious and
would take much too long to describe in such an account as the
present.
The sexes differ so much in the adult state that they would
perhaps hardly be recognised by the uninitiated as belonging to
the same species. The males are much more slender and of
smaller dimensions than the females, and they do not retain the
marvellous leaf-like appearance which characterises the latter.
The length of the male varies from 60 to 70 mm. The head is
somewhat quadrangular, generally of the same colour as the
remainder of the body, and possesses two or three red ocelli
which are situated between and slightly behind the level of the
bases of the antenne. The eyes are rather globular and very
prominent. The antennee—which are placed between the eyes—
are pilose and very long, often attaining 32 mm.; they are
composed of twenty-four joints which are smaller near the head,
becoming gradually larger in the middle and finally small again
at the extremities. When in motion the insect carries the
antenne forward either in a horizontal or semi-vertical position,
but when at rest they are placed backwards as shown in the
figure.
The prothorax is heart-shaped and tapers a little toward the
posterior end. The mesothorax is broader than the prothorax
and carries the tegmina, which are 13 mm. long and of parchment-
like consistence. The latter—which only cover a small portion
of the wings—are green but frequently spotted with brown or
red. The metathorax is of about the same width as the mesc-
thorax and bears two large membranous wings which extend
when folded almost to the posterior end of the body. The wings
are of delicate texture and are furnished with a regular network
of nervures and nervules;, in repose they fold up like a fan.
They are marked in several places with smaJl dark red streaks.
The first three abdominal segments become successively broader
until the maximum width of 24 mm. is reached in the fourth
abdominal. At each side of this segment there is a circular
transparent spot surrounded by a brown ring, and from here
the body becomes gradually narrower, finally terminating in a
bifid protuberance. The abdomen is exceedingly flat, and the
110 MR. H. 8. LEIGH ON THE [Feb. 2,
laminated expansions are generally pale green on the dorsal and
rich emerald-green on the ventral side. The colour, however,
varies considerably in different individuals, some being darker
green than others, and I have seen one or two quite yellow
specimens. The legs with the exception of the prothoracic
femora are entirely brown, and being prettily decorated with
many shades of yellow look rather like small pieces of decaying
leaves. ‘The expansions of the prothoracic femora are of a green
and brown colour; they are elegant in shape and, compared with
those of the female, are very small.
The females are mucb larger and more unwieldy creatures
than the males. They attain a length of from 95 to 100 mm.
The head is large; not quite so quadrangular as in the male and
is devoid of the red ocelli. The eyes are much less prominent
than in the male,and the antenne are very short, being composed
of only nine segments.
The prothorax takes the form of a shield with a slight furrow
in the centre. The mesothorax becomes much broader than
the prothorax and bears the remarkable leaf-like tegmina. The
latter, which generally remain in a state of repose, are of a most
remarkable structure and remind one of leaves to their minutest
detail. They are large and ample, being 54 mm. in length, and
generally cover the greater part of the abdomen. Each tegmen
is similar to half a leaf and has on its internal side a large
nervure from which at intervals smaller nervures emanate ;
these in turn eiving off subordinate veins which form a complete
ramification throughout the whole structure. The leaf-like:
form of the tegmina is carried still further since they are
adorned with many irregular rust-coloured markings similar to:
those often found on leaves.
The metathorax is broader than the mesothorax and attains.
at its posterior end a breadth of 18 mm. The hind wings, which
are so conspicuous in the male, are represented in the female by
two very small processes, covered by the tegmina and entirely
hidden from view.
As in the male, the first three abdominal segments become:
broader in regular sequence until the maximum width of about
44 mm. is reached in the fourth abdominal segment; from this.
there is a gradual tapering of the segments to the posterior end
which terminates in a bifid protuberance. The abdomen of the
female, although possessing flat expansions of considerable size,
is very bulky compared with that of the male, and even increases.
in volume as the ova mature. The general colour of the abdomen
is green of varying tints, and it is almost impossible to describe
one colour as applicable to all or even to many individuals.
Perhaps pale leafy-green is the commonest colour of the dorsal
side whilst the ventral is rich emerald-green. St. Quintin says.
he has “bred some entirely yellow and several of a crushed-
strawberry colour, while a few were of an amber-brown.”
Similar spots to those which were found on each side of the
1909. ] LIFE-HISTORY OF THE LEAF=INSECTe DE
fourth abdominal segments of the male occur also in the female,
but instead of being transparent they are often opaque and of
a brownish-red. colour. The first three abdominal segments are
generally ornamented at their edges with brown markings, and
there is frequently a brown patch at each side of the seventh
abdominal segment. Further, there is often an interrupted
brown mediodorsal line extending from the sixth abdominal
segment to the anal extremity.
The femora of the prothoracic legs are very large and foliaceous,.
having flat expansions which are green with many brown and
yellow markings. The mesothoracic legs are also of moderate
size, and on the posterior lobe of each femur there is a large
and conspicuous brown spot. All the iegs, however, have these
laminated expansions to a greater or less degree and the general
appearance of a female is that of a leaf.
Becquerel and Brongniart (1894) have carried out spectroscopic
investigations on the colouring-matter of the Phylliums with
a view to ascertaining whether it presented the same optical
characters as the chlorophyll of leaves. They found that the
spectrum of the Phylliums scarcely differs from the spectrum
observed through living leaves, but slight distinctions were seen
when compared with solutions of chlorophyll.
The majority of the males assumed the imago state long before
the females. The first male appeared on March 8th, 1908, and
others followed at irregular intervals until May and Thora, when
the maximum number was reached. From this time there was
a gradual diminution in the number of males. The first female
emerged on May 22nd, 1908. They continued to appear until
the end of August. Morton (1903) speaks of his first male as
appearing aborts six weeks before the ae female, and says that
nearly all the males were dead when this individual appeared.
St. Quintin (1907) also notes that his first male emerged one month
before the first female. I do not know whether the majority
of the males arrive at maturity before the females in the
Seychelles, but from the foregoing evidence such a state of affairs
seems quite possible.
The males are very active and are quite capable of flying a
few yards. Their activity is increased after dark, when they
usually move briskly over the leaves and occasionally fly from
one bough to another. Being so agile they are rather difficult
to inuadlle. and if touched generally. “drop or take a short flight
to some neighbourhing object. The females, on the other hand,
are very sluggish insects and seldom move far. They are quite
unable to fy," and if from any accidental cause they lose thei
foothold, can only ease the fall by means of their tegmina
which spread out like a parachute.
The life of the male is of about four or five weeks duration
and food is taken as in the larval stages, The females are much
longer lived and survive eight or nine weeks at least; they eat
much more than the males and consume in this period an
112 ON: THE LIFE-HISTORY OF THE LEAF-INSECT. [ Feb. 2,
immense quantity of foliage. The male generally selects his
mate during the night, and copulation takes place at some time
between 8 a.m.and 3 p.m. the following day. The sexes are
united for at least two hours.
I have not been able to ascertain so far the exact number of
eggs produced by one individual, but think that from eighty-five
to a hundred is the probable number.
Summary.—The Phylliums are dependent upon a very warm
and moist atmosphere, and are therefore more or less confined
to the islands in the tropical zone: in all stages they are very
similar, both in colour and habits, to various plant-structures.
Post-embryonic development is slow and takes place by a gradual
increase in size of the individual, adults only differing externally
from the young larve in the possession of fully developed
tegmina and wings. The sexual dimorphism is pronounced, the
females being large and foliaceous, whilst the males are much
smaller and although flat are not characterised by such a leaf-
like appearance as the females.
Tn conclusion, I wish to express my indebtedness to Professor
Hickson and Mr. Hewitt for their kind assistance.
REFERENCES.
1749. Rose v. Rosennor, A. J. Insekten Belustigung. II.
Theil: Orthopt. pl. 17, figs. 4, 5.
1767. Lixnzus, Carotus. Systema Nature. Ed. xii, tom. i. pt. 1.
1787-1815. Srotn, Caspar.—Representation exactement colorée
daprés nature des Spectres ou Phasmes, des Mantes, &c.
Amsterdam.
1839. SeRVILLE, AUDINET. Histoire naturelle des Insectes
Orthopteres. Paris.
1843. Gray, Groree R. Description of several species of the
Genus Phylliwm. Zoologist, vol. i. pp. 117-123.
1856. Murray, Anprew. ‘Notice of the Leaf-Insect lately
bred in the Roy. Botan. Garden of Edinburgh.” Edinb.
New Philos. Journ. vol. 111. pp. 96-111.
1859. Westwoop, J.O. Catalogue of the Orthopterous Insects
in the Collection of the British Museum—Phasmide.
Part I. London.
1871. Jory, Dr. N. “Contribution a Vhistoire naturelle et a
Yanatomie de la Mouche-Feuille des iles Seychelles.”
Mém. Acad. Sci. Toulouse (7) tom. 11. pp. 1-28, 3 pls.
1883. Dr Borre. Compt. Rend. Soc. Ent. Belgique, vol. xxvii.
p. exh.
1887. Bronentar?, CHartes. ‘ Note sur le développement de
la Mouche-Feuille de Java (Ph. siccifolium).” Ann. Soe.
Ent. France (6) tom. vil. Bull. pp. bexxiv—Ixxxvil.
1890. Hennecuy, L. F. ‘“ Note sur la structure de l’enveloppe
de Vceuf des Phyllies.” Bull. Soc. Philom. Paris (8)
vol. 11. p. 18.
1909. ] ON THE MAMMALS OF MATABELELAND, 113
1894. BecquEreL, H., er Bronenrart, Cu. “ La matiére verte
chez les Phyllies.” Compt. Rend. Acad. Sci. Paris,
vol. exviil. pp. 1299-1303.
1898. SHarp, Davrp. Account of the Phasmide, with Notes
on the Eggs. Willey, Zoolog. Results, Cambridge,
pp. 49-94, pls. vii.ix.
1903. Morton, W. “ Notes sur l’élevage des Phyllies.” Bull.
Soc. Vaudoise, xxxix. pp. 401-408, pl.
1904. Kirpy, W. F. Synonymice Catalogue of Orthoptera.
British Museum. Vol. i.
1907. St. Quintin, W.H. “Leaf-Insects in Captivity.” Ento-
mologist, vol. xl. pp. 73-75, and 147, pl. iv.
1908. Sr. Quintin, W. H. “Notes on the Life-history of the
Leaf-Insect.” Naturalist, no. 618, pp. 235-238, pls.
XXV., XXXVI.
EXPLANATION OF PLATE XXVIII.
Phyllium crurifolium.
The figures represent the adult male and female of the natural size,
and were photographed from living examples.
1. Female imago.
2. Male imago.
2. The Mammals of Matabeleland. By E. C. Causs, F.Z.S.,
Acting Curator of the Rhodesia Museum.
[Received December 29, 1908. ]
The Rhodesia Museum has acquired during the past two years
sufficient material to give a good idea of the mammalian fauna of
the Western Province of Southern Rhodesia, and it is upon this
collection that the present paper is based. For the sake of com-
pleteness, however, I have included the mounted examples of big
game that were in the Museum prior to this period, and also
a few species known to occur in the country, though up to the
present unrepresented in the Museum.
The only previous paper dealing with the mammals of this
area is that of De Winton ™*, being a list of a collection made by
Selous at Hssexvale. The species therein recorded are also
incorporated in this list.
Two interesting Bats, Rhinolophus empusa and Cleotis percivali,
are now recorded for the first time from South Africa south of
the Zambezi, hitherto being known only from the type localities,
Nyasaland and British Hast Africa respectively.
The probability of the different geological formations supporting
distinct vertebrate faunas was pointed out to me some time ago
by my former colleague, Mr. F', P. Mennell, and indeed this seems
to be borne out in a remarkable way by at least one group among
* P. Z.S, 1896, p. 798.
Proc. Zoou. Soc.—1909, No. VIII. 8
114 MR. E. C. CHUBB ON THE [ Feb. 2,
the mammals, viz. the Elephant-Shrews, one of which, Hlephantulus
rupestris myurus, occurs Wherever there is an outcrop of granite,
while the other, Vasilio brachyrhynchus, is found only on the
schists. Although separated generically on dental characters,
these two shrews are very closely allied in other respects and are of
very similar habits. They are never found living together, how-
ever, for whereas the latter is common at Bulawayo on the schist,
yet on the granite only two miles west of the town, and also at
Helenvale about 15 miles north, and the Matopos some 40 miles
south, it is replaced by Hlephantulus rupesiris myurus.
As a general rule the mammals here get their new annual
coats at the beginning of the rainy season, about October or
November, and the contrast between the old faded coat and the
new one is very marked in jackals killed in, say, August and
November.
The country slopes down from a line running north-east
through Bulawayo, at an altitude of about 4500 feet above the
sea, to the Zambezi and Limpopo Rivers on either side, which
are about 2000 feet and 1500 feet respectively.
I must express my indebtedness to Mr. Oldfield Thomas,
Dr. Knud Andersen, and Mr. R. C. Wroughton, who have been
good enough to compare and identify a number of specimens
for me.
The native names are those of the Matabele, an offshoot of the
great Zulu nation, who trekked up from the south about seventy
years ago. The letters ‘‘c” and “q” in these names represent
clicks. “c” is mnade by pressing the tongue against the front
upper teeth and quickly withdrawing it, as isance. It is much
the same sound as the expression of vexation “tut.” “q” is
made by curling the tip of the tongue upwards against the roof
of the mouth and quickly withdrawing it, as igaga. This click
sounds like the ‘ pop” of a cork when extracted from a bottle.
1. CERCOPITHECUS PYGERYTHRUS Cuv.
There are two flat skins, without skulls, of this Monkey in the
collection. One was obtained near Inyati and the other in the
Matopo Hills, where the species is fairly common.
Native name: /nkau.
2. PAPIO sp.
G. lv Oct, 190%, —Matopos:
2. 6 Sept., 1907. es
Native names: Jndwangu and Imfene.
3. GALAGO MOSSAMBICUS Pet.
36. 9 May, 1908. Bulawayo.
Sul 3 Aue, 908: a5
S. 19 March, 1908. Gambo’s, Manzamnyama River.
Nocturnal. It lives in a thorn-tree (Acacia horrida), and
sleeps during the day in the hollow trunk or in a roughly
1909.] MAMMALS OF MATABELELAND. TLS
constructed nest of grass and twigs ina fork among the branches.
Its food consists chiefly of insects, which it catches with its hands,
but the gum of the tree on which it lives is also relished. This
it takes in its hands and licks.
Tts eyes are a rich dark red or mahogany colour, with a round
pupil.
Native names: Ugwintu and Umpuguryone.
4. RHINOLOPHUS DARLINGE K. And.
6 2. Mazeppa Mine, Gwanda. In al.
Native name for all Bats: Ululuwane.
5. RHINOLOPHUS AUGUR ZAMBESIENSIS K. And.
329. 5 April, 1907. Bulawayo.
&& @, 8 Ayoall, Mos, 5
3d, 2. Mazeppa Mine, Gwanda. In al.
Lives in caves in granite kopjes and also in mines.
6. RHINOLOPHUS EMPUSA KK. And.
3. Mazeppa Mine, Gwanda. In al.
This is interesting as being the second known individual
example of the species, and also the first record of its occurrence
south of the Zambezi. It was described on a specimen from
Zomba, Nyasaland *.
7. RHINOLOPHUS HILDEBRANDTI Pet.
3 Q. 22Sept., 1907. Malute Reef, Pandangwe, Gwanda.
4 $g,5 2. Mazeppa Mine, Gwanda. In al.
Lives in mines.
8. HrpposIDERUS CAFFER Sund.
@. 12 Oct., 1907. Mazeppa Mine, Gwanda.
23,49. Mazeppa Mine, Gwanda. In al.
9, CL@oTIS PERCIVALI Thos.
2 imm. Mazeppa Mine, Gwanda. In al.
3,29. 18 Aug., 1908. Mazeppa Mine, Gwanda. In al.
10. NycreriIs CAPENSIS Smith.
S$. 12 June, 1907. Bulawayo.
36,32. 15 June, 1907. Bulawayo.
3,52. Sept. 1907. Mazeppa Mine, Gwanda. In al.
Four out of the five females in alcohol are pregnant.
11. VESPERTILIO MINUTUS Temm.
3g. 25 June, 1908. Bulawayo.
6. 5 duly, 1908. Fs
36,32. July, 1908. Bulawayo. Im al.
* Ann. Mag, N. H. (7) xiv. p. 378, 1904.
g*
116 MR. E. C. CHUBB ON THE [Feb. 2,
This Bat makes its appearance just after sunset. It flies
rather low, and frequents in considerable numbers the neigh-
bourhood of cowsheds for the insects which abound over manure-
heaps.
12. ScoroPHILUS NIGRITA Schreb.
$6. 5 June, 1908. Bulawayo.
13. MINIOPTERUS SCHREIBERSI Natt.
g. 1 March, 1907, Bulawayo. In al,
©. 26 Aug., 1907. 4
©. 17 Aug., 1907. Gravesend Farm, Bembezi River.
This species often flies into houses in the evening.
14. ELEPHANTULUS RUPESTRIS MYURUS Thos. & Schw.
@. 24 Sept., 1907. Syringa.
236, 2. 20 April,1908. Mt. Silozi, Matopos, 4400 ft.
@. 7 May, 1908. Near Bulawayo.
June, 1908. Helenvale Farm, near Bulawayo. (In alcohol.)
Diurnal. It lives under boulders on granite kopjes or among
the heaps of large stones which frequently occur on the granite
formation. Its food consists of various kinds of insects—the
stomach of one that I found was full of leaf-cntting ants.
Though shy, venturing only with great caution away from its
retreat, and returning at the slightest approach of danger, it is
also inquisitive, and will sit under the ledge of a rock eyeing one
with curiosity and apparent unconcern for its safety.
Average measurements, taken in the flesh, of the three Silozi
examples :—Head and body 121 mm.; tail 133; hind foot 35;
ear 31; proboscis 14.
Native name: Utswebe ulukulu.
15. NasiLico BRACHYRHYNCHUS Smith.
3. 6Sept., 1907. Bulawayo.
OQ, 5B Seyi, WO
©, 31 July, 1908. a3
This species appears to be confined to the schist formation, and
its colour harmonizes well with the red soil. Like the former
species, it lives among heaps of stones. The example killed on
5th September contained one well-developed fcetus. Two were
shot, while the third was caught in a trap baited with fish.
Average measurements, taken in the flesh, of the three speci-
mens :—Head and body 108 mm.; tail 104; hind foot 29; ear 20.
Native name: Utswebe ulukulu.
poe)
16. CRocIDURA sp.
©. 20 June, 1908. Bulawayo.
2. Bulawayo. In al.
Native name: Utswebe.
1909. ] MAMMALS OF MATABELELAND. ay
17. Fevis reo Linn.
5. 9 Oct., 1907. Near Eagle Vulture Mine, Gwanda.
The Lion is frequently met with in various parts of the country.
Native names: /stlwane, Impanwu, Ingwonyama.
18. Fevis pPARDUs Linn.
The Leopard is by no means uncommon in many parts of the
country. It lives in the Matopo Hills, and I know of a number
having been killed recently at Belingwe.
Native name: J/ngwe.
19. FELIS sERVAL Erxl.
Kahlele’s, Matopos.
June, 1908. Gwaai River.
Native name: Jndhlozi.
20. FELIS OCREATA Gmel.
6. 1 Oct., 1907. Near Bulawayo.
Native name: J/gola.
21. FeLis caRacaL Gild.
18 June, 1906. Near Bulawayo.
Native name: /ntwane.
22. CYNHLURUS JUBATUS Erxl.
The Cheetah is occasionally killed in the country, and I know
of one which was shot recently at Essexvale.
Native name: Jhlozi.
23. VIVERRA CIVETTA Linn.
9. June 1907. Kahlele’s, nr. Mt. Silozi, Matopos.
Native name: /simbaluti.
94, GENETTA LUDIA Thos. & Schw.
16 March, 1908. Bembezi Diamond Fields.
g. 25 Sept., 1908. Bulawayo.
Native name: /nsimba.
25. GENETTA RUBIGINOSA Puch.
2. Kahlele’s, Matopos.
Native name: /nsimba.
26. Munees cAFER Gmel.
A skin without skull; killed at Wankies in September 1908.
Native name: Uwobo.
27. MuNGOS GAUUI Smith.
$6. 1 July, 1907. Inyamandhlovu.
©. 17 July, 1907. Bulawayo.
118 MR. EB, C. CHUBB ON THE [Feb. 2,
g. 19 July, 1907. Bulawayo.
6. 19 Sept., 1907. Syringa.
@. 23 Sept., 1907. 55
Diurnal and common. It may be seen crossing roads or foot-
paths at all times during the day.
Native names: Uducakidi and Uwobo.
28. MuNGos ALBICAUDA Cuv.
6. 4Sept., 1908. Syringa.
This specimen has a black tail, as is often the case in this
species. Said to live in kopjes and to feed on rats and mice. Ivis
grey.
Native name: /duhwa elimnyama.
29, HELOGALE PARVULA, Sund.
6 @. 28 Oct., 1907. Near Bulawayo.
May 1907. Lukozi River, Wankies Dist.
Diurnal. It lives in colonies among the boulders of granite
kopjes, and feeds on insects.
Native name: Ubutietwetwe.
30. CynicriIs SELoUSI de Wint.
@. 23 May, 1907. Bulawayo.
©, 21 June, 1907. 5
QO, ih Dalby WOO .
2. 1 July, 1907. Inyamandhiovu.
@. 95 Aug.,1907. Bulawayo.
Qs Sess WO , “5
és Is Ame, U0 ~
O-.
27 June, 1908. Near Bulawayo.
Nocturnal and common. It lives in holes in the ground, and
feeds on small mammals and birds.
Native name: /duhava.
31. PROTELES CRISTATUS Sparrm.
A native skin from 16 miles south of Bulawayo.
Native name: /sance.
32. Hya@na crocuta Erxl.
¢o. June 1908. Gwaai River.
Not uncommon in the forest-belts, known as ‘ gusos,’” where
its white chalky droppings are often seen.
Native name: /mpisi.
33. CANIS MESOMELAS Ehrenb.
@ imm. 15 Jan., 1908. Bulawayo.
Common, chiefly nocturnal.
Native name: Lhanka.
1909. | MAMMALS OF MATABELELAND. 119
34. CANIS ADUSTUS Sund.
g. 15 Nov., 1907. Kana River.
Native name: /gowa.
35. OrocyoN MEGALOTIS Desm.
A head-skin of this species from near Wankies is in the
collection.
36. Lycaon protus Temm.
©. 1Sept., 1907. Sebakwe.
Antelope Mine, 8. of Matopos (skull only).
Native name: /yanyana.
"37. AONYX CAPENSIS Schinz.
1 Dec., 1907. Belingwe.
Native name: /ntini.
38. MELLIVORA RATEL Sparrm.
Belingwe.
Native name: Umantswane and Ulinda.
39, IcroNYX CAPENSIS Kaup.
3 April, 1907. Bulawayo.
6 2 imm. 8 Nov., 1907. Bulawayo.
3. 22 May, 1908. Bulawayo.
dg. 16 July, 1908. Near Bulawayo.
Nocturnal, and not uncommon.
Native name: /qaqa.
40. FuNISCIURUS CEPAPI Smith.
6. Sept., 1907. Syringa.
6. 14 Oct., 1907. Bulawayo.
6. 1 Dee, 1907. Tokwe River.
632. 19 April, 1998. Mt. Selozi, Matopos.
This Squirrel has two very different habitats. It lives on
granite kopjes and also in the forests of mopane (Copaifera
mopane).
Native names: Ubuhlula, Isikale, and Uwodo.
41. GRAPHIURUS PLATYOPS Thos.
17 Oct., 1907. Syringa.
21! Oct., 1907. ey
OQ, Os
°
42. GRAPHIURUS GRISELDA Schwann.
1 April, 1908. Bulawayo.
16 Sept., 1908. fs
+0 +O
120 MR. E. C. CHUBB ON THE [ Feb. 2,
43, TATERA LOBENGUL® de Wint.
Gerbillus leucogaster de Wint. P. Z. 8. 1896, p. 806.
g. 10 June, 1907. Springvale Farm.
2 9.11 S8ept.,1907. Bulawayo.
S. 23 Sept., 1907. Buck’s Reef, Gwanda.
©, 23 Oct., 1907, Kbami River.
©. 16 Nov., 1907.” Kana River.
p. 23 Nov., 1907. Swena’s, Gwamayaya River.
Essexvale (Selous).
Mammary formula: 2—2 = 8.
Lives in holes in the ground, and is partially diurnal.
Native name: /mbeba.
44, OTOMYS IRRORATUS AURATUS Wr.
3. 10 June, 1907. Springvale Farm.
Native name for all Rats: /gundwane.
45, DENDROMUS MESOMELAS Brants.
3. 24 May, 1908. Helenvale Farm, near Bulawayo.
Caught in the nest, which was formed of woven grass-blades
and suspended among tall grass about 4 ft. 6 in. from the ground.
It appears to be very like the English Harvest-mouse in habits.
46. Mus nigricauDA Thos.
9, 27 April, 1907. Bulawayo.
Oo, ff Ovi, 190n
©. 28 May, 1908.
Crepuscular, nocturnal, and strictly arboreal. It lives in the
common thorn-tree (Acacia horrida) and feeds upon the gum which
exudes from it. The first example was taken alive with four
young from a tree that was being cut down.
Mammary formula: 1—2 = 6.
99
99
47. Mus rarrus Linn.
EKssexvale (Selous).
‘Very common in houses near Bulawayo.
48. Mus cHrRySoPHILUS de Wint.
6 April, 1907. Bulawayo.
12 Oct., 1907. 5
10 June, 1907. Springvale Farm.
16 Sept., 1907. Syringa.
. 22 Sept., 1907. Pandangwe Kopjes, Gwanda.
@. 12 Nov., 1907. Indabambi’s, Shangani River.
Q. 24 May, 1908. Helenvale Farm.
Exssexvale (Selous).
Os O, +O Os Os OF 24
s
Mammary formula 1—2=6. This is our commonest rat. On
rough granite country and kopjes it takes grass into the crevices
between rocks to form nests. It occasionally enters houses.
1909. | MAMMALS OF MATABELELAND.
49. Mus Aaurtcomis de Wint.
BaZa Selsgo, p. 80.
Essexvale (Selous).
50. Mus cotonus Brants.
36,2. 10 June, 1907. Springvale Farm.
6. 4Sept., 1907. Bulawayo.
S. 18Sept., 1907. 5
2. 9Nov., 1907. Shangari River.
@. 12 Nov., 1907. Indabambi’s, Shangani River.
Very common, and enters houses.
51. Mus micropon Pet.
16 March, 1907. Bulawayo.
14 Feb., 1908. <5
8 April, 1908. 9
3 May, 1908. 5
20 April, 1908. Kahlele’s, Matopos.
25 May, 1908. Helenvale Farm.
This like Mus colonus often enters houses.
ee pene ee
52. Mus muscutus Linn.
@. 24 March, 1907. Bulawayo.
Very common.
53. LEGGADA MINUTOIDES Smith.
g. 7 Nov., 1907. Tjokos’, Shangani River.
54. SACCOSTOMUS HILD& Schwann.
25 Oct., 1907. Khami River,
3. @ Feb., 1908. Near Bulawayo.
This species has only been found on the granite formation.
55. AcoMys SELOousI de Wint.
IP, Zo So USO, jos tec
Essexvale (Selous).
56. ARVICANTHIS PUMILIO DILECTUS de Wint.
6 2. 10 June, 1907. Springvale Farm.
57. ARVICANTHIS DORSALIS Smith.
3 4 Sept., 1907. Bulawayo.
d 5 Sept., 1907. *
©. 18 Sept., 1907. 5
g. 21 Sept., 1907. Buck’s Reef, Gwanda.
18 Oct., 1907. Bulawayo.
3
Diurnal.
122 MR. E. C. CHUBB ON THE [ Feb. 2,
58, GeorycHus NIMRODI de Wint.
P. Z. 8. 1895, p. 808.
Essexvale (Selows).
59, PEDETES CAFER Pall.
¢. 13 June, 1907. Colleen Bawn Mine, Gwanda.
g. 18 Jan., 1908. Bulawayo.
@. 14 Oct., 1908. Bulawayo.
Native name: IZyelane.
60. THRYONOMYS SWINDERIANUS Temm.
©. 16 Sept., 1907. Bulawayo.
ye. 1 Aug., 1908. es
6. 9 Oct., 1907. Springvale Farm.
Mammary formula 2—1=6.
Native name: Jvondo.
OQ,
je)
61. Hysrrix sp.
The Porcupine occurs in the country, but 1 have not succeeded
in obtaining an example of it yet.
Native name: Jnungu.
62. LEPUS ZULUENSIS MICKLEMI Chubb.
6. 13 April, 1907. Bulawayo.
6. 15 April, 1907. 7
29.4 May, 1907. rr
Gan Usrdiulyy 19 Oe 50
6. 27 April, 1908. 3
6. 2 May, 1908. 3
@. A August, 1908. A
1 July, ISO Nyamandhlovu.
Very common.
Native name: Umvundhla.
63. PRONOLAGUS RUDDI RANDENSIS Jameson.
Syringa.
Confined to granite kopjes.
Native name: /nteletja.
64. DAMALISCUS LUNATUS Burch.
36 2. Near Mangwe.
Native name: /nkolome.
65. ConNocH#TES TAURINUS Burch.
6 2. Near Manewe.
Native name: /nkonkoni.
1909. | MAMMALS OF MATABELELAND. 128
66. CEPHALOPHUS GRiMMr Linn.
9. Near Manewe.
6. 9 July, 1907. Insiza.
346,22. 24 Oct., 1907. Mt. Silozi, Matopos.
3. Inyati (albino).
Native name: /mpunze.
67. OREOTRAGUS OREOTRAGUS Zimm.
36, 2,yg. Near Mangwe.
dS yg. 2. 24 Oct., 1908. Mt. Silozi, Matopos.
@. 17 April, 1908. Near World’s View, Matopos.
Native name: gogo.
68. RAapPHicerus campestris Thun.
36 9. Near Manegwe.
9. 4 June, 1907. Springvale Farm, near Bulawayo,
3. 25 Oct., 1907. Mt. Silozi, Matopos.
Native name: /nqina.
69. RAPHICERUS SHARPE! Thos.
6. Aug., 1908. Near Inyati.
3. Oct., 1908. Sinonombi.
Native names: Jsanempa and Lsanhlunywana
70. Kopus ELLIPSIPRYMNUS Ogilby.
9. Near Mangwe.
Ss. Oct., 1908 (head only). Sinonombi.
Native name: /sidumuha.
71. CeRVICAPRA ARUNDINUM Bodd.
6 2. 24 Oct., 1907. Mt. Sitozi, Matopos,
Native name: Umzigi.
72, ANPYCEROS MELAMPUS Licht.
26. Oct., 1908. Sinonombi.
Native name: /inpala.
73. Hipporracus EQuINuUS Desm.
6 2. Near Mangwe.
Native name: /taga.
74, HipporraGus NIGER Harris.
3 2%. Near Mangwe.
Native names: Umtjwayili and Ingwalati.
124 ON THE MAMMALS OF MATABELELAND, (Feb. 2,
75. SrREPSICEROS STREPSICEROS Pall.
6 2. Near Mangwe.
Native name: /mbalabala.
76. TAUROTRAGUS ORYX Pall.
$ @. Aug., 1907. Kana River (skeletons).
Native name: Jmpofw.
77. BUBALUS CAFFER Sparrm.
The Buffalo is found in Northern Matabeleland, and I know of
two males being shot there in October 1908.
Native name: Jnyati.
78. GIRAFFA CAPENSIS Less.
@. Near Mangwe.
Native name: Jntudhla.
79, HipPporporaAMUS AMPHIBIUS Linn.
There are many hippos still in the country, and some were shot
at Belingwe this year.
Native name: Jmvwbu.
80. PoraMOCHGRUS CH@ROPOTAMUS Ill.
3. July 1908. Matopos.
Native name: Jngulugundu.
81. PHAcOCH@RUS ®THIOPICUS Pall.
3. Oct. 1907. Khami R.
Fairly common all over the country.
Native name: Jnrgulubi.
82. Equus BURCHELLI CHAPMANI Layard.
2, Near Mangwe.
Native name: /dube.
83. DicERos BICORNIS Gmel.
So. Jan., 1907. Gwelo.
Native name: Umkombo.
84. PROCAVIA CAPENSIS Pall.
36 imm. 13 Sept., 1907. Syringa.
SQ. 23 Sept., 1907. Pandangwe Kopjes, Gwanda.
2. 20 Feb., 1908. Between Lochard Siding and Inyati.
1909. ] ON PATHOLOGICAL OBSERVATIONS DURING 1908. 125
3.17 April, 1908. Near World’s View, Matopos.
3. 19 April, 1908. Mt. Silozi, Matopos.
One of the skins from between Lochard Siding and Inyati has
a large white patch on the top of the head.
Native name: Jmbila.
85. PROCAVIA BRUCEI Gray.
d. 24 April, 1908. Kahlele’s, Matopos.
Trapped in tree by natives.
Native name: Jnbila.
86. ELEPHAS AFRICANUS Blum.
The Elephant is to be found in Northern Matabeleland, and I
saw fresh spoor of a herd on the Kana River in November, 1907.
Native names: /ndhlovu and Inkubu.
87. MANIS TEMMINCKI Smuts.
6. 22 April, 1907. Wankies.
Native name: J/nkaka.
88. ORYCTEROPUS AFER Pall.
3. Aug., 1907. Mapisa’s, near Figtree.
®. 24 Oct., 1908. Shiloh.
Native names: /sambane and Iwombela.
3. Report on Deaths which occurred in the Zoological
Gardens during 1908. By H. G. Promer, F.L.S.,
F.Z.8., Pathologist to the Society.
[Received February 1, 1909. |
On January 1, 1908, the number of animals in the Zoological
Gardens was 3190 and during the year there were 2418 new
arrivals, making a total for the year of 5608. 1737 animals have
died during the year, that is roughly 31 per cent., but if from
this number we subtract 769 animals which did not live for
six months after their arrival in the Gardens—that is, those
which had not got used to their new environment—the per-
centage of deaths is reduced to 17 per cent., which is probably a
much fairer number to take.
Of those which have died, 1089 have been examined, and in
41 of these no cause of death could be found. Of the remainder
129 died from injuries of various sorts, or were lost, or not sent
for examination, and 30 died from exhaustion due probably to
depressed vitality from cold or darkness, or from inability to get
or to take food.
126 DR. H. G. PLIMMER ON PATHOLOGICAL [ Feb. 2,
The Tables which follow show the facts ascertained in bare
outline, and following t them are some short notes on the most
impor swat points conmacrerl with these facts.
Table I. sets forth the causes of death in each of the three
great classes of animals. Under Reptiles are included batrachians
and fishes.
Taste I.—Analysis of 993 deaths.
DISEASE. ‘Manas. Brrps. REPTILES.
BERS St it i = 2 === |
1. Microbic or Parasitie Diseases. | | See Notes.
MM ibenrculosisucceesceereresa ee Caeeeceel 59 88 | Wy (eauilbs
INS RGONS) Gagucoadedoododeastscsde so09829ed008 ue Bain ae Bare 2.
IW GME TE ae ene RE Ace Asan aeltncaes SaEAG lui MesdAwes ILLS ee || arc ae 3h
I Que TEE Hiee aa ai mee eer nReen ms bcntdcoseadcan 3 18 il | 4
Trypanosomiasis Paik ecelacanceceuisn 1 20 We Has 3.
IEIERUOERCEEIEUNES, aoccancnccoossesosnboncn| ceased | ongeee 24. 6.
Pneumomiar econ ees tec ee 4 29 19 Wo
Septiceemia ub cian esata 12 AD accctarsits |
Pseudo- HUF IBIHOUULOMIS sussese-ouey/seatoonne Bis) AIT ere ie meses ds
TBGRGENEIGIS) ” onenao ateonbovosedasooosnszoan sul 1 eee ay lt ae oink
Worms Behe e eae pau hase ON, cer She ees 15
Itch.. geal ee eee wl eet
Pr otozoal infection és | ecb sl User cea IL 8.
Y. Diseases of Lungs.
Broncho-pneumonia ...s.............-| 44. Ieee ecm Vite sear |
Cloinexasinin Orr IMTAGHS co o-5ee5 650008 oa5o00 29 | 100 | 14 | @:
|3. Diseases of Heart. | |
| |
TREICRIRCINNIS) 4).oepbososedee cooecneadeacanall a 3 il
Hatty degemeratiom ...........02..020c02| cere Stabile sree
| }
|4. Diseases of Liver.
| Hepatitis ved Oo ea Aire Die eee, pl ae
| Fatty degeneration A en Roe 1 13 | 3
|5. Diseases of Alimentary Tract.
SS bometittisyaso-e2 ssn niavect > ete eee eeionl il DAM AG Re
Gastritisy fa .ca essere erases een 4 il 2
Gastric uleer ... 0.6.2... ILL 1 1 lO:
| GCREROLMUAUIS, Geoesos-oonsucenanccaase 23 2 27 ea
Winheritisnspeee ss ie caeteeec ans | 57 | 139 31 5)
(COMMS — csaseae odesepe eye ee ae LD pep ye leg bens bt: 1
Acute tympanites- a ae ha arenes alec lee ectees?
| Tmitussusce pioneer tents Eee enacee Dai Wee pre se
| OVONGRUIGHION Gooceorasadesoswndarcarvecnose|) uranpo A gS We cect a
| J PPS BUOTAINIHIS Socipoogounanendsacnooundesppag otc 15 Ciel ny cece
| 6. Diseases of the Urinary and
| Reproductive Organs. |
ANG phiritish, eee Meee eee terre | 4 TRAST GoM |
TOMS ENTTTMETON (ONE CVACILGH onooovccsenec.n)) aves 2 il |
|7. Various.
Senile changes ..... WONG |MeweE PR EST oh re Lei
Anemia without ascertainable eee 5
bo
oo
tH
bo
LI09e OBSERVATIONS DURING 1908. 127
Table IT. shows the distribution of these diseases amongst the
principal classes of Mammals.
TABLE TI.— Distribution of diseases amongst the Mammals.
Disease.
ee 5 |
Prinates.
Carnivora.
Rodentia.
Ungulata
a
“Rm yeROWIGSUS..54).55 ooovocnoecbanve se caycsnneal GS 3 | 14 2 | 5
ila ame eee are tei ctontict hei ccc mes 3 Peed ete, |
Trypanosomiasis ..00...cccsccceeeecseee eee Mesto wie de tis baba ac
I EMC DyaaVMDINE: Papen adace nda naS ude Aad sap Ren Anas i @ 14 6 2 6
epiiCocmilaerrene mecme neers al 6 12 5 is 4 1
Pseudo-tuberculosis ......cccccccecsceeee] we [Foe 3
FLV Gabi Sy: swrreeess inane eeeent uaa || wes cally aie
VV foren stp tere arte ceyan. tecaieotren ate gaa 2 x
Itch Pee Cee nal atte kan] 2 1 234
Broncho- -pheumonia EAaring saceosanicenees i 3) 3 10 2 if
| Comegasingrn Oi? WHEY ..ccsvesoeseonsceocc| 1B 5 5 3 3
| IRE TBERTROD ITTY che adsadaakedanconcdesaadedceters| heh wun. || fines) as 2 1
| Hepatitis ....... iI
| Patty degeneration of liver er
| Stomatitis Paste yoiasuibae oie esoesenert leat selet fs 1 Be di ah at
Gastritis 1 1 Il al
Gastric ulcer a 2 2 ee 4
Gastro-enteritis ee peas er an hg 8 3 3 AL eh
LDL HSERHIS a. so piecafeireeacoelsteteee tt dota ge tay. ple 11 9 IB po Bf ako
Colitis MEL cae neon eree toca Sia ees ve 2 | 2
Acute tympanites | dotead aanaiccen amnion Nee teat i eo: eo fi sdee eee at
TUREMESMISCAUHOM Gooorsoosnrccayeaacovoaveas| one wi sa iment
Peritonitis doh aa aden ot RaMR REE 1 il 3 iil 3
| INI) SOU PUSIEH atic dot nop ohasbapacitonmevnadbaacoul|\ Mun? 2 se Bap il
| Senile changes ........ Se oaeu to tceeoe ed Bad NS haan 1 FORA An ae
| Anemia (cause unknown) eNeferinds seunaa| Miidee 1 1 00 ey oe
| | |
Since the lesions of tuberculosis and mycosis are somewhat
similar in appearance, care has been taken to separate them
accurately, and the following Table sets forth their distribution
amongst the different classes of birds.
Tasre [I1. —Comparative Table of the incidence of Tuberculosis
and Mycosis 1 in the v various classes of bir ds.
|
Tuberculosis. Mycosis.
|
|
|
(Jt)
rag
We}
TERICISYSs Helsiee s dee aneinaaane pigibbabedod
IPH GinSh ie a baeceesoecas Rebosacoeuce| 2 5
TS vote aie eer aati 0 2 11 |
INTIS coacan ns eperoeenaoustobdcasc bon 4, |
(Cliliviralipa: tracreatedeetepscesss No. cool 12
bo
bo 6
(Cen llitnt teeny Maen Sacson ncdaretosnoet
filbairi Ceara. Sean eerrareeestee: ae 1 5
SHHPEUHOUCOIOVSS)) .ododoonosnauodcunsoanu0 2
128 DR. H. G. PLIMMER ON PATHOLOGICAL [ Feb. 2
The seasonal variations of the principal diseases are very much
as might be expected, and there seems to be little of importance
to record in this connection. Tubercle remains pretty constant
all the year round; mycosis has been more prevalent during the
second and fourth quarters ; pneumonia and broncho-pneumonia
were more prevalent during the first and fourth quarters; and
diseases of the alimentary canal have been more marked in the
latter half of the year.
There has fortunately been no epidemic amongst the animals
during the year, and disinfection has been carried out after every
case of infectious disease, so as to lessen, as far as possible, the
risks of infection. It will be noticed that there has not been a
single case of tumour, cancerous or otherwise, in the animals
examined during the year.
Notes on the foregoing Tables.
The following notes refer to a few points of special interest in
connection with the diseases mentioned in the Tables.
1. Since the beginning of last year all reptiles have been
systematically examined, and tubercle has been found in 17 cases
out of a total of 161; and in 5 snakes, tubercular deposits have
been found in the lungs. Pulmonary tubercle in reptiles has
not, I believe, been described before, and the first specimen
found has been preserved in the Museum of the College of Surgeons.
Tubercle found in reptiles has so far been all of the avian type,
both microscopically and in cultures,
The 88 cases of tubercle in birds seems a large number, but it
compares favourably with the number of cases recorded” as
occurring in the Berlin Zoological Gardens between December
1903 and August 1905. Between these dates 459 birds were
examined there, of which 118 were tuberculous; our figures show
that of 479 birds examined only 88 were tuberculous. In two
parrots examined the tubercle was of the human type, in all the
rest it was distinctly avian.
Of the 35 cases of tubercle in monkeys nearly all were of the
human variety, two, however, were of a distinctly bovine type,
and in one case the bacilli were of the avian type.
2. In four of the cases of mycosis the disease was due to
Aspergillus niger, the lesions being similar to those produced by
the Aspergillus fumigatus which was the organism associated
with the remaining 51 cases.
3. Under the general term Malaria I have grouped a series of
cases which, so far as I am at present able to state, are caused
by parasites belonging to either the group of Halteridium or of
Proteosoma. In these birds the parasites were certainly the
cause of death, but they were also found in 12 other birds which
died from other causes.
A. Several of the filaria found have not been described before
and are still under investigation. Im some cases two quite
different kinds of filaria have been found in the same host, as in
* WVirchow’s Archiy, vol. 90.
1909.] OBSERVATIONS DURING 1908. 129
two Lawes’ Birds of Paradise. The difficulty of identification is
much increased by the fact that the parent worms often cannot
be found. The filaria found in a Bearded Lizard is, I believe, the
first found in a reptile.
5. A hitherto undescribed trypanosome has been found in a
Francolin which died from mycosis, which is larger than any
described so far in birds.
6. Hzmogregarines have been found in 40 reptiles, and in 13 of
these, I believe, for the first time. In the cases in which death has
been attributed to them the anemia and blood destruction have
been so extreme as to make the diagnosis quite certain ; sometimes
50 or even 60 per cent. of the erythrocytes have been affected.
7. Pneumonia in reptiles is of two kinds—one a specific inflam-
mation of the lung, the other a traumatic inflammation due to the
irritation produced by ascaris eggs and embryos deposited there.
In some cases masses like tubercles are formed around the source
of irritation. 13 of these cases were of this irritative kind.
8. This protozoal infection occurred in a bull-frog, and the
lesions were identical with those described by Danielewsky in
1889 in his account of the only other case on record in an edible
frog. The causative organism which was present in the blood
and the exudations in this case was a Hexanvitus (really Octomitus).
9. Of the 29 mammals which died from congestion of lungs,
22 were suffering from rickets, 9 very severely.
10. It will be seen that gastric ulceration is more widely distri-
buted amongst mammals than is usually thought to be the case.
11. The large number of cases of enteritis has led to an
investigation into the probable causes. There appear to be
three different kinds of enteritis in the Gardens—one which is
caused by bacteria, one which is caused by protozoal organisms,
and a third variety probably due to errors in feeding. An effort
is being made to separate the various kinds, so that at any rate
those due to feeding may be got rid of. The very large per-
centage of cases of gastro-enteritis in the reptiles will be noticed ;
in 58 out of 161 cases death was due directly to this condition,
and in the remainder, which died of other diseases, the same
condition was present in varying degrees. It would seem worth
while to consider whether the present unnatural and unphysio-
logical method of feeding the snakes has any connection with the
very large incidence of inflammations of the alimentary tract
occurring in them.
12. In these 10 cases there was very profound anemia, with
considerable blood changes. J think that these cases were in all
probability parasitic in origin, but no cause, parasitic or other,
could be found.
Several probably undescribed intestinal worms have been found
during the year, and are still undergoing investigation.
The work condensed in this paper has been carried out in the
laboratories at the Gardens and at the Lister Institute.
Proc. Zoon. Soc.—1909, ING@ UBS 9
130 MR. C. TATE REGAN ON [ Feb. 16,
February 16th, 1909.
FREDERICK GILLETT, Esq., Vice-President,
in the Chair.
The Secretary read the following report on the additions made
to the Society's Menagerie during the month of January,
ie SMS)
The number of registered additions to the Society's Menagerie
during the month of January was79. Of these 54 were acquired
by presentation, 20 by purchase, 2 were received on deposit, and
3 were born in the Gardens,
The number of departures during the same period, by death
and removals, was 158.
Amongst the additions special attention may be directed to :—
One Desert Fox (Vulpes leucopus) and two Gmelin’s Sheep
({Ovis orientalis) § 2, the latter new to the Collection, from
Persia, presented by Capt. T. H. Keyes, F.Z.S., on Jan. 5th.
One Markhoor (Capra falconeri) 3 ,from N.W. India, pre-
sented by Lt.-Col. 8. H. Godfrey, C.I.E., on Jan. 18th.
One Red-cheeked Scimitar Babbler (Pomatorhinus erythrogenys)
and two Occipital Blue Pies (Urocissa occipitalis), from India,
purchased on Jan. 21st.
One Javan Drongo (Crypsirhina varians), from Java; one
Dumont’s Grackle (lino dumonti), from the Aru Islands; two
Black-throated Lorikeets (Zrichoglossus nigrigularis), two Green-
naped Lorikeets (Z'richoglossus cyanogrammus), and three Green-
winged Parakeets (Aprosmictus chloropterus), from New Guinea—
the Drongo, the Black-throated Lorikeets, and the Green-winged
Parakeets being new to the Collection,—purchased on Jan. 18th.
Four Black-cheeked Love-birds (Agapornis nigrigenis), from
N.W. Rhodesia, new to the Collection, purchased on Jan. 15th
and 16th.
Mr. C. Tate Regan, M.A., F.Z.S., exhibited sketches illustrating
changes in coloration of some Fishes in the New York Aquarium,
and made the following remarks :—
“In September 1907, I paid several visits to the New York
Aquarium; the director, Mr. C. H. Townsend, very kindly spent
a good deal of time in showing me the fishes and calling my
attention to many interesting details. I was particularly struck
by the colour-changes in some of the tropical Sea-Perches from
the Bermudas, and I made some notes on the coloration of four
species, Viz. :—
“1. Hpinephelus striatus.—l devoted some hours in all to the
study of this species. The normal coloration, if such a term can
be used, may be described thus: ground-colour greyish-olive
with irregular paler spots ; body crossed by about five broad dark
brownish bars; a black spot on the upper part of the bar which
1909. | COLOUR-CHANGES IN FISHES. 131
encircles the caudal peduncle. This description would probably
apply to some, and usually to most of the specimens observed at
any given time, but they were constantly changing, and I came to
the conclusion that there must be several different systems which
could work independently, either in harmony with or in opposition
to each other. These systems were (1) the ground-colour except
(2) the pale spots ; (3) the upper and (4) the lower halves of the
-cross-bars and (5) the caudal spot. If these systems worked in
harmony the fish would be uniformly coloured, varying from
almost white through various shades of greyish-olive and brownish
to black. Working independently, the cross-bars might become
defined and become even quite black while the ground-colour,
remained nearly white, or if the ground-colour became darker the
pale spots might or might not appear. The caudal spot was often
fading out and suddenly reappearing. Certain changes could only
take place in a definite order ; the lower halves of the cross-bars
never appeared before or persisted later than the upper halves ;
the caudal spot was never paler than the bar on which it was
placed, although it could entirely disappear if the bar had faded
out; similarly the bars were never paler than the ground-colour,
and although the pale spots could become quite black with the
rest of the ground-colour, they could not darken independently
of it. All the changes mentioned above could take place either
slowly or very rapidly.
“2. Bodianus punctatus.—So recently as 1902 Prof. Evermann *
distinguished between L. fulvus, with the ground-colour lemon-
yellow, B. ruber, with the ground-colour bright scarlet, and
B. punctatus, with the ground-colour brownish or blackish-olive.
He wrote that these had usually been regarded as subspecies, but
that until the fact of intergradation was established, it was best
to treat them as distinct species. Individual specimens in the
New York Aquarium are continually changing from one so-called
species into another. In this species three distinct cclour areas,
viz. the upper, middle, and lower parts of the fish, separated by
straight longitudinal lines, may be recognised. I observed a
uniformly yellow fish suddenly assume an orange tint on the back ;
this gradually deepened into a brownish-red whilst the sides and
lower parts became paler. A dark greenish fish changed suddenly
into a red fish, and a uniformly red fish became pale on the sides,
the upper and lower parts remaining unchanged. A. fish with the
back chocolate-coloured became darker above and paler below, so
that the nearly black back was sharply defined from the almost
white sides and lower parts.
“3. Mycteroperca bowersi.—A beautiful fish of a brilliant red or
crimson, with black edges to the fins. I was only able to watch
this species for a few minutes, but saw one red fish become pale
pink and another dark olive-green; another in the tank was
brownish, and in one the black edges of the fins had disappeared.
* Bull. U.S. Fish. Comm. xx. pt. 1, pp. 149-150.
Q*
132 DR. F. WOOD JONES ON THE [ Feb. 16,
“4, Hemulon flavolineatum.—Ground-colour changing from
pale yellow to deep bronze, with numerous undulating stripes of a
blue of varying intensity. Two blackish longitudinal stripes, the
lower ending in a spot at the base of the caudal fin, suddenly
appeared and disappeared. Similarly, a few irregular broad
blackish cross-bars could be turned on and off.”
Mr. E. G. B. Meade-Waldo, F.Z.S., read extracts from a letter
he had received from Dr. Einar Lonnberg, C.M.Z.8., on the
hunting of the Sea-Elephant on South Georgia, and called attention
to the necessity of steps being taken to prevent its extermination.
The following papers were read :-—
1. The Fauna of the Cocos-Keeling Atoli, collected by
F. Wood Jones. By F. Woop Jones, B.Se., F.Z.S.,
with the assistance of other Authors.
[Received December 3, 1908. |
(Text-figures 7-9.)
The fauna of the Cocos-Keeling group has been several times:
investigated, but it has never been thoroughly worked out. The
species enumerated in the following lists were collected during a
stay of fifteen months in 1905 and 1906, and the collection, of
most orders, may fairly be considered as complete.
Darwin visited the group in 1836 and stayed for only ten days,
from April the 2nd to the 12th. Wallace in his ‘Island Lite”
quotes Darwin’s list of the fauna (p. 275).
Dr. H. O. Forbes visited the islands in 1879, arriving on
January 18th, and staying till February 9th, and he amplified
considerably the list made by Darwin; unfortunately his
collections were lost in returning to Java, and so the additional
species that he observed have not been specifically recorded (eae
Naturalist’s Wanderings in the Eastern Archipelago,’ 1885).
On the 20th of August 1885, Mr. W. E. Birch, on behalf of
the Straits Government, landed and made inquiries about the
islands and their inhabitants; with him, as naturalist, went the
Rev. E. C. Spicer. The expedition visited most of the islets, and
remained for eight days in the atoll, but in the report (Straits
Blue Book 1885) no light is thrown on the condition of the
fauna.
In succeeding Blue Books are scattered notes, made by the
Commissioners, on some of the most striking features of the atoll
fauna, but most of this information is mere interpretation of local
legend, and is of no value.
Dr. H, B. Guppy came to the islands in 1888, and has written
1909. |] FAUNA OF COCOS-KEELING ATOLL. 133
an excellent account of their physical features in the Scottish
Geographical Magazine (vol. v. 1889, pp. 281, 457 and 569), but
he has not, so far as I am aware, published any description of the
fauna.
Since the atoll lies 12 degrees south of the Equator, it is of
necessity subjected to some seasonal variations, and though these
are greatly moderated by the influence of the South-East trade
winds, they are sufticiently pronounced to affect the time of
appearance of most of the insect fauna. It is therefore obvious,
that a stay of a few days could not produce, even with the most
industrious collecting, at all a representative series of the insect
fauna. The present list need not therefore be taken as indicating
that a great increase has taken place in the fauna within the past
thirty years, for it is more properly an index of more protracted
collecting at all seasons of the year’s cycle.
The atoll consists of some two dozen typical ‘“ low ” islands, that
rise but a few feet above mean tide-level, and enclose a lagoon
some 8 miles in diameter. The whole group liesat 12°10°S., 96°
52° E.; and is separated by upwards of six hundred miles from
Java, and upwards of five hundred from Christmas Island, its
nearest neighbour.
In 1827 the atoll was settled by the pioneer member of the
Clunies-Ross family, and ever since that time the destinies of the
group have been swayed by three generations of the same family.
The islands were uninhabited before the advent of the Clunies-
Ross dynasty. In 1901 a Telegraph-station was erected on Pulu
tikus, for the working of the cable between Perth (Australia)
and Mauritius.
The South-East trades blow for some three hundred days each
year, and the climate is subject to no sudden changes save infre-
quent cyclones; the mean temperature is 81° Fahr., and the
annual rainfall averages something over 70 inches.
MAMMALIA.
The Rats of the atoll are of some interest, apart from their
economical importance. When the original settlers arrived in the
group in 1827, rats were already established on one of the islands,
and this island was named Pulu tikus, or rat island, in con-
sequence.
From 1827 till 1878 these rats remained as the only repre-
sentatives of their family, and continued to be almost confined to
the island of their settlement. Towards the end of 1878 a ship,
‘The Robert Portner, was wrecked in the atoll, and left as her
legacy a flourishing colony of Mus decwmanus ; since then other
wrecks, and trade intercourse, have added Mus rattus to the island
fauna. The products of these separate invasions have remained
distinct until to-day, and they have fairly sharply defined areas:
of distribution. The “original rat,” as it is called, remains
confined to Pulu tikus. MZws decumanus is most abundant on
134 DR, F. WOOD JONES ON THE [ Feb. 16,
Pulu Selma, and some of the smaller islands to the south. d/us
rattus has its stronghold on Pulu gangsa, where it is very
abundant, but it also occurs on the other islands in company with
M. decumanus. When Dr. H. O. Forbes visited the atoll in 1879,
he noted the greatnumber of rats—these were the ‘ Robert Portner
rats’ that had invaded the islands in the previous year, and had
multiplied to an extraordinary extent. He makes no note of the
black rats, and does not mention the race of ‘ original rats’ on
Pulu tikus.
During Darwin’s visit the only rat in the islands was this Pulu
tikus race, and of it he says:—‘‘ These rats are considered by
Mr. Waterhouse as identical with the English kind, but they are
smaller and more brightly coloured” (‘ Naturalist’s Voyage,’ p.461);
and Wallace (‘Island Life,’ p. 275) adds, ‘‘ We have here an illus-
tration of how soon a difference of race is established under a
constant and uniform difference of conditions.”
In the seventy years that have passed since Mr. Waterhouse
described the rat, the difference appears to have become more
marked, and a description of the race is justified for the reason
that the modern introduction of rat virus into the atoll may
easily exterminate it.
Haternal appearance.—The rat isa slenderly built and sleek
looking animal]; its general colour is a warm russet-brown. The
fur of the back is coarser, and some hairs stand out that are almost
black, and are 30 mm. in length; these long dark hairs are more
numerous over the hind end of the body than over the shoulders.
The general colour of the body is a rather rich brown, the belly
being but little lighter than the back in most specimens. The
shorter hairs are grey at the base and bright red-brown at the
tips; the longer hairs are dark brown to black. The fore limbs.
are somewhat lighter than the rest of the body, but the hind
limbs are of the general rich red-brown. The under surface is
only slightly lighter than the back, and isa light warm brown,
not grey; the chin and throat are somewhat lighter. The under
surface of the scrotum of the male is covered with long bright
red-brown hairs, brighter than those of any other part of the
body ; the distal extremity of the scrotum is bald, the naked skin
being a dark purplish brown.
The vibrisse—many of them more than 60 mm. long— are
numerous and dark-coloured. The ears are almost naked, they
are oval and prominent and average 22 by 15 mm. The hind
feet are large and long, their soles are dark, almost purple,
coloured; the digits are slender, the claws are long and much
curved. The tail very slightly exceeds the length of the head and
body, it is darker than the body colour; it is finely scaled, 12 rows.
to the centimetre, and sprinkled with numerous fine black hairs.
There are two pairs of pectoral mamme, and three inguinal pairs,
2-3=10. The tibie are almost straight, having practically no.
‘““bow ” forwards.
The Skull.—The skull is long and narrow, and is delicately
1909. } FAUNA OF COCOS-KEELING ATOLL. 135
built. The nasals extend posteriorly to the anterior margin of the
orbit, are 15°5 mm. long, and 2°8 mm. wide at their broadest
part. Basal length of skull 35-38 mm.; greatest breadth 18 mm..
Interorbital constriction, least breadth 6 mm. Interparietal,
length 5 mm., breadth 10 mm. Length of base of anterior root
of zygoma 6 mm. Palate, length 20 mm.; breadth, outside m*
9 mm., inside m’4 mm. Palatine foramina 8 mm., extending
1 mm. behind the anterior edge of m'—this is a constant feature.
Alveoli (back of incisors to m') 12 mm. Upper molar series.
6°5 mm. Incisors pale yellow in the lower jaw, orange in the
upper jaw.
The measurements in millimetres of a series of specimens are
as follows :—
|
Head ; |
and Tail. me | ae | von Forearm.) Sex. |
| Body. oot. | (tibia). oot. |
i} | |
1 190 120M) MELONI BO 20 See a
2 207 268 40 AS eile 20 33 4
3 195 200 40 50) 20 SOPe ers
4 195 | 191 38 42 22 25 a
5 205 | 215 40 52 19 34 e
6 1S 185 40 50 18 32 3
7 192 | 216 40 | 50 19 34 ef
8 200 | 205 40 45 20 30 Q
9 180 | 185 AOne iss 20 3U fe)
10 163 | 158 39 AD 22 29 Q
11 150 155 38 39 22 27 e)
12 147 149 36 385) belies 24, Q
13 215 230 42 45 | 22 36 Q
14 198 | 202 39 AS We SX) 32 2
TSW ISZ 1-490 39 Asis | is 32 Q
peneen || 270 160 SaWey Aoi galas 20 38 fe)
lice gle 120 130 a He BB aye ale/ 32 2
| }
Habits.—The rat is very abundant on Pulu tikus, and although
it has been steadily trapped, and hunted by dogs, for the last four
years, it does not appear to diminish in numbers. As many as
forty will still be taken in wire traps beneath the Telegraph
quarters in a night, and outside the little area of the station the rat
swarms everywhere. It is by no means a house rat, and although
it is common in and around the houses it is still more abundant
in the outlying parts of the island. It is splendidly adapted to
its environment; it has altogether given up burrowing, and it.
lives in the piles of coconut husks and in the crowns of the palms.
It is certain that, at times, the young are born at the tops of the
palms, which are 60 to 70 feet high. The rats ascend the trees with
the greatest ease, and they run from one palm to the next, across:
the interlacing fronds, very much after the fashion of the squirrel.
They are by no means strictly nocturnal, doubtless from their
long freedom from any enemies, and they are to be seen running
about at all times of the day. They feed almost entirely upon
the fallen coconuts, and all the nuts that I have ever seen opened
136 DR. F. WOOD JONES ON THE [ Feb. 16,
have been gnawed by rats, and not torn open by Birgus latro, as
they are often said to be. Even those that are trapped around
the Telegraph quarters are almost invariably full of coconut, and
most come into the houses to obtain water rather than for food.
Before the Telegraph-station was built no water was available,
save that collected during showers in the bases of the palm-leaves.
It has been killed in vast numbers since the opening of the
Telegraph-station on Pulu tikus in 1901, but its numbers do not
seem to have diminished at all. Considering its extreme
abundance it does very little harm, save eating the copra stored
on the island, and it has never adopted the practice of nibbling
the green nut, that makes the rats on the other islands such an
economical curse.
Mus decumanus, typ.—This is the pest of the atoll, and does
incalculable damage to the coconut plantations. A perpetual war
is waged against it by means of traps and dogs; natives are
specially told off for the purpose of rat killing, and all the dead
rats are regularly recorded. Apart from eating copra, and
damaging stores, it has learned to climb the palms, and destroys
countless nuts by nibbling them through, just where the stalk joins
the husk. It is known in the islands as the ‘ New’ or ‘ Norwegian’
rat, and is always distinguished by the natives from the Pulu
tikus rat.
Mus rattus, typ—Is common on Pulu gangsa and some of the
small southern islets, and I have had one specimen from Pulu
Selma, where it is not nearly so common as J. decumanus. It
does not exist in such numbers as does J/. decuwmanus, but it has
learned the same destructive habits. It is never met with on
Pulu tikus.
Mus musculus Linn.—By no means abundant, although for a
few months in 1905 it became very common in Pulu tikus; on
that island it does not appear to have been seen in any numbers
until about 1903, and it was doubtless introduced with stores from
Singapore.
A herd of feral deer lived for long in Pulu luar. The animals
were introduced from Java and from Singapore, and consisted of
two species, the Sambar (Cervus hippelaphus) and the Kedang
(Cervus muntjac). All were dead when I left the islands in 1906,
and the only trace of their presence was a well-marked “ browse
limit” to the trees, a novel feature in a coral-island landscape.
Waifs and strays include bats of some small species that did
not appear during my stay in the atoll, and that are said to be
the Pipistrellus murrayi Andrews, from Christmas Island. <A
large monkey is also said to have been washed ashore, and to have
long survived in the islands; the method of its advent is not
known.
The domesticated animals include dogs, cats, and pigs; sheep
have been turned down, with but little success; and rabbits
liberated on Pulu luar do not seem to have multiplied greatly.
1909. | FAUNA OF COCOS-KEELING ATOLL. 137
IOVIDS:
CoLUBBIFORMES.
(1) CARPOPHAGA WHARTONI Sharpe.
Native name, ‘ Pergam.”
- Jntroduced from Christmas Island, but now (1906) practically
extinct in the atoll.
RALLIFORMES.
(2) RaLLus PHILIPPENsIs Linn.
Native name, ‘“* Ayam utan.”
Very abundant on all the islands, and is everywhere very tame,
it being a matter of some difficulty to make it take wing. It
feeds on the shore when the tide is out, but it may also be seen
perched high in papaia trees eating the ripe fruit, and it has a
bad name for eating the eggs of domestic fowls. It nests in
September, in tufts of grass, about a foot from the ground ; it
lays from two to six eggs, very like the English Corncrake’s. The
young are all black when hatched, and can run directly they are
out of the egg. The call-note isa shrill grating sound, and in
the breeding-season the cock adds a deep croak not unlike the
noise made by frogs. This species is not found in Christmas
Island.
LARIFORMES,
(3) SrerNA FULIGINOSA Gmel.
Native name, “ Burung dali.”
Breeds in Keeling Island but not in very great numbers. It
keeps very much to itself on the breeding grounds, and lays one
egg in the sand above the beach rise, on the western side of the
island. By sailors it is called the whale-bird. Although not
uncommon, and often seen at sea, far from the atoll, it does not
oceur on Christmas Island.
(4) Anous stotipus Linn.
Native name, ‘“ Burung krok.”
Not resident in the Southern atoll, but a frequent visitor to the
lagoon. In Keeling atollon it breeds in great numbers. In June
the breeding-season has almost come to an end. The nests are
made about a foot above the ground, on little collections of sea-
tossed wrack. In the atollon all the ‘“noddies” build close
together in a rather limited area. Only one egg was found in
each nest.
(5) Gyais canpipA Wagler, Isis, 1832, p. 1223.
Native name, ‘‘ Burung chuit-chuit ” (onomatopeic).
Darwin noted the bird in 1836 (‘ Naturalist’s Voyage,’ p. 462),
and H. O. Forbes gave an account of it in 1879 (‘ Naturalist’s
Wanderings,’ p. 34). The bird agrees with the description of
138 DR. F. WOOD JONES ON THE [ Feb. 16,
Wagler :—‘TIrides nigro-ceerulee: rostrum nigrum, basi coeru-
leum ; lingua sublata rostro brevior ; pedes cerulei, palma alba,
ungues nigri.”
In a good description of this bird (Cassin, U.S. Explor. Exped.
p. 389) the feet are said to be “pale blue, having a deeply
indented yellow membrane.” In Gould’s ‘ Birds of Australia,’
vol. vii., the feet are described as orange, and are figured of that
colour in the accompanying plate. In other accounts it is stated
that the feet are yellow or brown. The bird, as I have seen it in
the Cocos-Keeling group, invariably has the feet entirely blue,
the web being slightly lighter than the toes.
Text-fig. 7.
Egg of Gygis candida, laid on the branch of a grongong tree (Cordia
subcordata).
The bird is a common one, and occurs in all the islands. It
breeds in the Southern group as well as in Keeling Island.
Although the bird feeds mostly on fish, it may often be seen
hovering around the papaia-trees and eating the soft fruit. It is
quite arboreal in its habits, and its one egg is laid sometimes at a
height of 40 feet from the ground. Most of the eggs were found in
September, but on visiting Keeling Island in June two eggs were
found after very little searching. The egg is almost perfectly oval
(40-43 mm. in its long axis, 21-22 mm. in its short axis); it is
1909.] FAUNA OF COCOS-KEELING ATOLL, 139
cream-coloured, mottled and streaked with olive-brown. It is
laid on the branch of a tree (Pisonia inermis, Cordia subcordata,
Guettarda speciosa) wherever a slight irregularity in the bark
suffices to hold it steady (see text-fig. 7). In all the cases that I
have seen the egg was laid in the long axis of the branch, and the
parent bird sits across the branch in the process of incubation.
From the fact that three nesting sites were used twice over, it
would seem as though the suitable branches for balancing the egg
were somewhat limited. The birds take it in turn to sit, or rather
stand, on the egg, and they invariably leave it by falling backwards
Text-fig. 8.
Gygis candida sitting on its egg.
This is the same ege as is shown in text-fig. 7.
off the branch, in order not to disturb it with their feet; great
caution is taken in getting into position on the egg again (see text-
fig.8). Incubation lasts 36 days; the newly hatched young is buff-
brown, and it does not move from the tree until it has assumed
its white feathers. A bird hatched on Sept. 3rd, 1905, did not
move from the site of its hatching till Oct. 22nd; it then moved
higher up the branch, andremained, with slight changes of position,
till Nov. 12th, when it took its first flight (see text-fig. 9, p. 142).
Many eggs were watched and in no case did any accident befall
them, but a young bird, hatched on Noy. 17th, fell from its insecure
140 DR, F. WOOD JONES ON THE [ Feb. 16,
position on the fourth day, and was killed. This bird does not
occur on Christmas Island, and is not seen far from the atoll—it is,
in fact, a sign that the atoll is near when one is seen on the
voyage from Java.
ARDEIFORMES,
(6) Demrecrerra sacra Sharpe.
Native name, ‘ Blakok.”
Exists in fair numbers, especially in Pulu atas, where as many
as twenty may be seen fishing together. Birds in the white
phase and in the grey phase mix freely together; though
it is said in the islands that white pair with white, and
grey with grey. ‘The nest is made in the Pisonia trees, and two
pale blue eggs are laid. I obtained specimens in January.
In Pulu atas I have also seen a solitary individual of a larger
species of Heron, that, in the distance, appears to be pure white ;
this individual was well known to the natives of the atoll.
PELECANIFORMES.
Suborder FREGATI.
(7) Frecara Aquiia Linn.
Native name, “ Burung itam.”
Does not breed in the main atoll, but nests in large numbers
in Keeling Island. The nest is a slight collection of twigs, stolen
from the nesting Gannets, and placed on the flat tops of the
Pemphis bushes. ‘The nests are crowded together in thousands ;
one egg only is laid, which is pure white and its surface is shining ;
eggs were abundant in June. The bird is so tame in Keeling
that it has to be driven from the egg, and those birds that are
sitting in the hot sun, with thei wings half spread, may easily:
be approached and lifted from the ground.
(8) FReGATA ARIEL Gould.
This species is more common than the last. Its habits are
similar, and it nests freely with the larger species. It is also
called Burung itam by the natives.
Suborder Sun &,
(9) Suna sua Linn,
Native name, “‘ Burung bebek.”
I have never seen this bird in the main atoll, and its numbers
are somewhat limited in Keeling Island. It lays one or two
dirty white eggs on the bare ground, at the top of the western
sand-beaches.
(10) Suna apsorrr Ridgw.
Native name, “ Burung gangsa.”
Is not common in Keeling, and, so far as I have observed, does
1909.) FAUNA OF COCOS-KEELING ATOLL, 141
not come to the Southern group. The face is black, and the bird is
considerably larger than the other two species. One or two eggs
are laid on the bare ground. The black mask gives the sitting
bird a very curious appearance, and it leaves its egg only after
having, with great solemnity, vomited large quantities of fish ;
it makes no attempt to resist being pushed from the egg, and
when the egg is taken, and the fish has been vomited, it quietly
walks away. No skin was obtained.
(11) Suna piscArrix Linn.
Native name, ‘ Burung main main.”
The commonest of the three species, and generally called
at Booby it by s sailors in these seas. Exists in very great numbers,
and is seen in all the many age changes of plumage, from a
uniform light brown to white. It lays one egg, in a nest placed,
as a rule, high up in trees; but numbers build on Pemphis
bushes in Keeling. The birds commonly perch on trees, and are
often taken from the rigging of ships.
Suborder PH AETHONTES,
(12) Puarrnon rupricaupa Bodd.
Native name, “ Burung buntut.”
Does not exist in great numbers, but still breeds in Pulu atas.
This species, as well as P. fulvus, is to be seen everywhere
between the atoll and Java Heads.
PASSERIFORMES,
Fam. PLOCEIDA.
Mr. H. O. Forbes mentions Ploceus hypoxanthus as one of the
nesting species of the atoll; but no examples of it exist to-day,
(13) Munia oryzivora Bp.
Native name, “ Burung glatek.”
Introduced, and has multiplied greatly ; it is now one of the
commonest birds of the atoll.
Fam. TuRDIDA,
(14) Meruna ERytHROPLEURA Lister,
Introduced from Christmas Island, and now very common,
especially on Pulu tikus. The bird is remarkably tame, coming
freely into rooms to obtain water. Has a rather pleasing song,
which it utters in September and October. The nest is built in
November, and the egg is very like that of the European Redwing.
Fam. ZOSTEROPID 4x
(15) Zosrrrors NavTAuis Lister.
Introduced from Christmas Island. Confined to Pulu luar,
where it exists In some numbers. In Christmas Island it is
142 DR. F. WOOD JONES ON THE [ Feb. 16,
known as Burung Waringin, but in Cocos-Keeling it has earned
the name of Burung chinta or “ love-bird.”
“‘ Golden plover,” native name “ Burung blebis,” and “ Curlew,”
are said to breed on some of the islands, but I never found the
egos, nor did I ever obtain a bird for identification. A flock of
ducks is resident on Pulu panjang, and on the same island I have
seen many small waders, but the wise policy of the islands is that
these birds shall not be interfered with, and I cannot name the
species. I have, however, shot a stray Snipe (Scolopax gallinago
Linn.) from a small flock on Pulu tikus, and on the same island
obtained a Swallow, Hiruwndo gutturalis Scop. ‘These are the only
visitors that came to the islands in my stay of fifteen months, and
the atoll would seem to be altogether out of the track of any,
save very wind-blown, migrants.*
Text-fig. 9.
Young Gygis candida, soon about to take its first flight ; the plumage is
still mottled with buff-coloured feathers.
* In North Keeling atollon are large warrens, the nesting sites of “ Mutton
Birds” ; during my visit to the atollon these birds were not present, but they
evidently visit the place, for the purpose of breeding, in very large numbers.
In all probability they are Puffinus brevicaudus or P. sphenurus.
1909.] FAUNA OF COCOS-KEELING ATOLL. 143
REPTILIA.
Species determined by G. A. Boutencsr, F.R.S., V.P.Z.S.
(A) LacerRTILtia.
Fam. GECKONID&.
(1) LeprpopactyLus Lucustris D. & B.
Native name, ‘“ Cheechak.”
A very abundant species, inhabiting dwelling houses, and also
living beneath the loose bark of coconut palms. Six or seven
eggs are laid at a time—as a rule beneath some convenient chink
of bark. It has the characteristic voice of its family, which has
given rise to its name.
(2) GEHYRA MUTILATA Wiegm.
Not nearly so common as the first species, and lives exclusively
out of doors, being always found beneath the bark of the coconut
palms. Jt is at once distinguished from No. 1 by its flattened
tail. It lays from three to four eggs, and the incubation period
is as long as two months (June to August).
(B) OrpHipra.
Fam. TYPHLOPIDE.
(3) TypHuops Braminus Daad.
Native name, ‘“ Ular minyah.”
This is not a common species in the islands, and its habits lead
to its being very rarely seen. It is most commonly found beneath
large coral boulders, and it is extremely active. It is found on
practically all the islands in the atoll. The average length is 150
millimetres.
None of these species is found in Christmas Island.
(C) CHELONIA.
Two species of Turtles are frequently speared in the lagoon.
They are Chelone imbricata iu., native name “ Pinnew sisih,” and
C. mydas L., native name ‘‘ Pinnew betul.” Both have ceased to use
the main atoll as a breeding place, for the native demand for
them is too great; but on Keeling atollon their nests are
common on the sandy beaches.
Accidental visitors.—From time to time other reptiles have been
washed ashore. Several large snakes have floated to the islands,
and have been picked up dead, or dying, on the beaches. Some
few have survived the journey, but none has made a successful
footing. At least two crocodiles have survived the six hundred
miles of ocean travel : one was shot by the Governor, and one, after
being repeatedly seen, disappeared after a cyclone.
144 DR. F. WOOD JONES ON THE [Feb. 16,
PISCES.
The fish of the group were not collected, but no doubt a rich
harvest awaits the investigator of the myriad smaller coral-
haunting species. The native names of those that are of the
most economical importance only are given here; and no
specific determination is possible in most cases.
(1) Ikan babi = pig fish, Balistes sp. (2) Ikan buntal =
inflated fish, Vetrodon patoca. (3) Ikan buntal besagi = square-
shaped inflated fish, Zetrodon sp. (4) Ikan blanah. (5, 6) Ikan
bandang and Ikan bandang laut. (7) Ikan buntut burih =
spotted tail. (8) Ikan chuchut, generic name for the numerous
Sharks. (9) Ikan dongol, Scarus sp., grows to a great size.
(10) Ikan grapu. (11) Ikan iju = green-fish, Psewdoscarus sp.
(12) Ikan jengot = bearded fish. (13) Ikan jengot karang
(karang = coral). (14) Ikan kakap. (15) Ikan kakap kuning =
yellow kakap. (16) Ikan kakatua iju, Searws sp. (17) Ikan
kakatua merah = red kakatua,alsoa Scarws. (18) Ikan merah =
red fish. (19) Ikan malam = night fish. (20) Ikan menyrat.
(21) Ikan padang doeh, the albicore. (22) Ikan p’dang, Histio-
phorus gladius. (23) Ikan pareh, Dicerobatis eregoodoo, grows
to a great size, being upwards of 13 feet across the back.
(24, 25) Ikan peteh, and Ikan peteh kuning. (26) Ikan puti =
white fish. (27) Ikan palo. (28) Ikansambar. (29) Ikan samsi.
(30) Ikan skagnol. (31) Ikan talam talam. (32) Ikan talang.
(33) Ikan tangiri, the king fish. (84) Ikan todak, the baracouta.
(35) Ikan trompet.
It is to be hoped that, with the free communication that the
Telegraph-station affords, some attempt may be made to get
together a collection of the fish of the atoll: and for the purpose
of assisting any collector, this incomplete list of the native names
is published.
LEPIDOPTERA.
LEPIDOPTERA RHOPALOCERA.
(Native family name, “ Kupu kupu.”)
Species determined by Mr. F. A. Hrron.
(1) Precis vinuipa Fabr.
This is the commonest butterfly in the atoll. It is most
adundant in July and September, and again in March and April.
The larva is blaek, and is covered with spines: food-plant is
Asystasia coromandeliana. The pupa is suspended by the tail
from the branches of its food-plant, it is mottled brown and has
no metallic markings. The pupal stage lasts for a fortnight or
three weeks.
(2) Hypotymnas Botrna Linn.
1909. ] FAUNA OF COCOS-KEELING ATOLL. 145
(3) Hypontmnas BoLina form NERINA Fabr.
An uncommon species, on the wing in April and August.
Specimens are in perfect condition, and the species is evidently
resident.
(4) Hypotimnas misrppus Linn.
A common species on all the islands of the group. One worn
specimen was taken on Christmas Island by Dr. Andrews. The
males and females of this species are very rarely seen together ;
the males flying high in the shade of the thiekly growing coco-
palms, and the female being generally seen flying low over the
herbage in open spaces. It is abundant from July to October,
and again from the end of February to April. The larva is
brown and spiny, the head is reddish and bears two spines.
Tt feeds on a succulent weed that grows all over the open
spaces in the atoll. The pupa is brown and has no metallic
markings ; it issuspended by the tail. Six days are passed in the
pupal stage. The female mimics Danais (Limnas) chrysippus
petilia Stoll, and is nearly always seen flying in company with
that species, whose habits of fluttering over the herbage it has
closely imitated. The male, on the other hand, flies strongly and
is not often seen fluttering near to the ground,
(5) VANESSA KERSHAWI McCoy. (Australian.)
Only four examples of this species were met with in fifteen
months, and since all were observed between the 15th and 17th of
May 1906, it is probable that they were waifs. Two rather worn
specimens only were captured on Pulu tikus.
(6) Danais (Lonas) cHRysrppus PEritiA Stoll. (Australian
& Christmas Id.)
Common on Pulu luar and Pulu tikus, but not so often met
with on the other islands. The larva is pale green, banded with
yellow and black. The food-plant 1s Asclepias curassavica. The
pupa is suspended by the tail, and is pale green or buff with small
golden dots. The average stay in the pupa is only 135 hours.
LEPIDOPTERA HETEROCERA.
Species determined by Sir Grorce F. Hampson, Bt., F.Z.S.
ARCTIADZA.
(1) Urernesa PULCHELLOIDES Hmpsn.
A very abundant insect on all the islands. It mostly frequents
the seaward side, for there the food-plant, Tournefortia argentea
Linn., lives most luxuriantly. The perfect insect, and the larve,
are to be seen at all times of the year, and season appears to make
no difference to its numbers.
Proc. Zoou. Soc.—1909, No. X. 10
146 DR. F. WOOD JONES ON THE [ Feb. 16,
Nocruipa.
(2) CutoripEA opsoLera Fabr.
No specimens were taken during 1905, and the insect only
became at all plentiful in June 1906.
(3) CHLORIDEA AssuLTA Guen.
Appeared at the same time as the last species, and was not
seen in any numbers.
(4) Crrpuis LoREY! Dup.
Not an abundant insect, and strictly confined to a seasonal
appearance. Taken in June 1905 and not again until the same
month in 1906.
(5) PropEnta LirroRALs Boisd.
Abundant on all the islands, on the wing most plentifully in
May and June. The larva feeds on a multitude of low-growing
plants. This very widely distributed species is found on Christmas
Island.
(6) SpopopreRA MAURITIA Boisd.
Several specimens were taken in June 1905, but in June 1906
it did not appear again.
(7) LeocymaA sERIcEArA Hmpsn.
Not abundant, appears in June and again in November. The
larva is pale green and bears a few scattered hairs, it feeds on the
leaves of the Waroo (Hibiscus tiliaceus Linn.). Not on Christmas
Island, though its genus is represented by Z. tebialis Fabr.
(8) Opuiusa coronata Fabr.
Only one specimen taken, and a few more seen, in June 1905.
Occurs on Christmas Island.
(9) OpHrusA MELICERTA Dru.
This is one of the most plentiful of the atoll insects, being
found wherever the bushes of Pemphis acidula (native name “ Kayu
burung”) are growing. It is on the wing practically all the year
round, but is most common from June to September. The larva
bears a wonderful resemblance to the twigs of the Pemphis acidula
or the Ricinus on which it feeds. It has a habit of dropping
from its twig when disturbed, but it always manages to get a
fresh hold of a lower twig, on the way down, although it appears
to be falling to the earth. It pupates in a few leaves spun
together at the ends of the twigs; the pupa is covered with a fine
bloom. Fourteen days are passed in the pupal state.
1909.] FAUNA OF COCOS-KEELING ATOLL. 147
(10) Remicia FRuGALIS Fabr.
Most abundant in May, and a few stragglers again in Sep-
tember. It flies by day, and is not uncommon in the grassy
spaces, where the undergrowth is kept cleared.
(11) Prusia cHatycres Esp.
The development of a garden on Pulu tikus appeared to be the
cause of the abundance of this species; but since the garden
plants were introduced as seed, there appears but little chance of
their having been the agent for its introduction. It swarmed in
June 1906, and the green larva was abundant on peas, tomatoes,
and almost every plant that was grown in the garden, The pupal
stage lasts only a week.
(12) Hyprna stricara Fabr,
One example only taken,
(13) CarEprRia sp.
This is a handsome insect, boldly marked with black and white.
It appears on the wing in September. The larva feeds on the
leaves of Pemphis acidula, in company with Ophiusa melicerta,
to which larvee it bears a considerable rough resemblance. It is
mottled grey on the back, and the belly is velvety black; at the
anal extremity is an upwardly directed tubercle with a bifid tip.
A collar of magenta colour surrounds the larva behind the head,
but in all ordinary attitudes this is hidden by a skin-fold. It
pupates among the little leaves of its food-plant ; the pupa has no
bloom, and the pupal stage lasts for twenty days. I never saw an
example of the perfect insect, save those that | reared from larve ;
and, so far as I could find out, no one on the island was acquainted
with the moth. The larve were by no means uncommon on a
few bushes in Pulu tikus, but I did not meet with them on any
other islands.
SPHINGIDE.
(14) Macroeiossa PAssALusS Dru.
By no means common, and more frequently seen as a larva than
as an imago. On the wing in September. Larva green with
fine bright pink side-stripes; feeds on Morinda citrifolia Linn.,
native name ‘ Mungkoodoo.” The pupa is a mottled yellowish-
brown.
(15) CePpHonopEs HyLAs Linn.
Common on Pulu tikus, but not often seen on the southern
islands. Frequents the flowers of the Papaia by day, and is
evidently the source of the error that bees are common in the
islets. When hovering in front of the food-plant, in the act of
laying its eggs on the leaves, the long hairs of the extremity of
the body stand out like a fan. Most abundant in June and
September, but stragglers may be taken in almost any month.
Os
148 DR. F. WOOD JONES ON THE [Feb. 16,
Larva feeds on the leaves of Guettarda speciosa Linn., native
name “ Melati.” When first hatched they are quite black, and they
only become green in their penultimate and ultimate skins.
They are subject to endless variation, and if bred in the dark,
very striking larve, coloured yellow and black, can be produced ;
the imagines resulting from these larve are normal. This insect
occurs on Christmas Island.
(16) Herse convoryuti Linn.
Not a very common insect. Found most plentifully on Pulu
Selma. The larva is either green or brown, and it feeds on
Convolvulus parviflorus Vahl. In the atoll this is the moth
specially named “(Rama rama,” although in the Straits that word
is the general name for all moths.
(17) CHa#RocAMPA VIGIL Guer.
The perfect insect is rarely seen, but the larva is very abundant
in September. It feeds on the leaves of Pisonia inermis (native
name “ Ampol”); it is green, but in its last skin it frequently
becomes putty-coloured. When fed in the dark the last skin is
almost invariably brown. A month is passed in the pupa. This
species occurs on Christmas Island.
GEOMETRIDAE.
(18) CHLOROCLYSTIS TENUILINEA Warr.
Not uncommon in June and August. The larva is green, with
darker markings, and is slightly hairy. Feeds on the sticky
flowers of Pisonia, and pupates among them.
PYRALIDA.
(19) Pyrauis MANIHOTALIS Guer.
Not at all plentiful.
(20) MELISSOBLAPTES sp.
Very abundant on all the islands of the group.
(21) ZINCKERNIA FASCIALIS Cram.
Always common. ‘This widely distributed species also occurs on
Christmas Island.
(22) MARASMIA VENIHALIS Walk,
One of the commonest insects. Appears in June, and is always
to be found about the station lamps.
(23) CrocrDOLOMIA BINOTALIS Zell.
Only a few examples met with in June.
(24) PacHyzANCLA LICARSISALIS Walk.
Abundant from June to August. The larva is green, with
1909. | FAUNA OF COCOS-KEELING ATOLL. 149
minute black spots ; it feeds on the leaves and flowers of Dicliptera
burmanni Nees. It pupates in the spun-up leaves, and the pupal
stage lasts for fourteen days.
(25) PacHYZANCLA STULTALIS Walk.
Abundant in June on all the islands.
(26) GLYPHODES INDICA Saund.
Not abundant.
PrEROPHORIDA.
(27) A brownish coloured Plume is very common in all the
islands. The larva feeds on Boerhavia diffusa W., which grows
everywhere in the atoll.
TINEINA.
(28) The solitary species collected was a very common one:
there are probably at least two others that are distinct.
There are therefore practically thirty species of Heterocera on
the Cocos-Keeling atoll, and there are more than twice as many
species in the fauna of Christmas Island; yet only five, very
widely distributed species, are common to the two places.
HYMENOPTERA.
Species determined by the late Col. C. T. Bineuam, F.ZS.
FORMICID 2.
(Native family name, “ Samut.”)
(1) OponromacHuUS H#MATODES Linn. %.
(2) PLAGIOLEPIS LonerPEs Jerdon. @.
Unfortunately this does not represent the whole of the species
of ants found on the atoll. The natives distinguish at least three
species :—(1) “Samut saman”; (2) “Samut arpi”; (3) “‘Samut
alus.” The other representatives of the family are unfortunately
mislaid.
FOSSORES.
(3) Srizus sp. (very near reversus Smith), Q.
(4) Pison Hospes Smith. ¢.
(5) NoroGoNIA SUBTESSELLATA Smith. ¢ Q.
None of the Fossores is at all numerous, and only very careful
collecting produced the very few specimens obtained in the course
of fifteen months.
None of these species is represented in the fauna of Christmas
Island.
150 DR. F. WOOD JONES ON THE [ Feb. 16,
DIPTERA.
Species determined by E. E. Ausren, F.Z.S.
ASILID&.
(1) PurLopicus sAvANUS Wied.
A common insect of very voracious habits. It is generally seen
sitting on the coral boulders, in the hottest sunshine. It hes in
wait for smaller diptera, and it will attack and kill even butter-
flies as large as Precis villida.
BoMBYLID.
(2) AnrHRAX sp. Only a very few examples met with.
DoLicHOPODID#.
(3) New genus and species (No. I.), near Psilopus.
(4) New genus and species (No. II.), near Psdlopus.
Both of these are common insects, but owing to their green
colour and small size, they are not at all conspicuous. They are
shining green flies that are most commonly seen running on the
leaves of the Hibiscus. They are extremely active, and are
generally found during the hottest hours of the day.
Muscip2.
(5) SarcopHaGa sp. No. I.
(6) SarcopHaca sp. No, IT.
Both are common species, frequenting the dead bodies of rats.
(7) Ruria TEsTAcEA Rob.-Desv.
Few examples seen, mostly upon the herbage of the seaward
side of the islands.
(8) PycnosoMA FLAvICEPS Macq.
The commonest fly in the atoll.
(9) OPHYRA CHALCOGASTER Wied.
MIcCROPEZID#.
(10) Nerrus LINEoLATUS Wied.
HIpPoposcip®.
(11) PsevDoLFERSIA sPINIFERA Leach.
Usually swarming over the plumage of the Frigate-birds. On
1909. ] FAUNA OF COCOS-KEELING ATOLL. 15]
Keeling atollon, where these birds breed in thousands, the fly is
often seen crawling about the nesting bushes.
CULIGID.
(12) The Mosquito of the islands is a species of Stegomyia, and
it exists in great numbers.
COLEOPTERA. i
Species determined by C. O. WarEruoust, P.E.S.; C. J. GAHAN,
M.A., F.E.S.; and G. J. Arrow, F.E.S.
® BRACHELYTRA.
(1) AtEocHARA sp. This genus is not represented on Christmas
Island.
CLAVICORNIA.
(2) DERMEsTES FELINUS Fabr.
A common species, found mostly in the dead bodies of rats.
Universal distribution, and common on Christmas Island.
(3) CoccINELLA TRANSVERSALIS Fabr.
A very abundant insect; not present on Christmas Island.
(4) TROCHOIDEUS DESJARDINSI Guer.
Not present on Christmas Island.
LAMELLICORNIA.
(5) ONTHOPHAGUS sp.
Genus not represented on Christmas Island.
(6) ANOMALA sp.
An abundant species that lives on the iron-wood (Cordia sub-
cordata). Freely attracted to light and most commonly found in
the lamps of the Telegraph-station. Not found on Christmas
Island.
(7) Poratra ACUMINATA.
The largest beetle found on the atoll. It is by no means
common, and not more than a dozen specimens were taken in
fifteen months. Not present on Christmas Island.
MALACODERMATA.
(8) CoryNETES RUFIPES De Geer.
Native name, ‘‘ Kutu copra.”
The most important insect of the atoll, from an economical
point of view. Called by Europeans the “‘ Copra beetle.” Exists
152 DR. F. WOOD JONES ON THE [ Feb. 16,
in myriads near to the sheds where the copra is stored. It will
swarm to anything oily, and was killed in thousands on the
refrigerator engines. It is very active, it flies by day, and has a
habit of creeping beneath the clothing and biting rather sharply.
It does not oceur on Christmas Island.
SERRICORNIA.
(9) MEGAPENTHES sp.
Not a common insect. It does not occur on Christmas island,
where MW, andrewsi Waterh. is the representative of the genus.
(10) MELANOXANTHUS MELANOCEPHALUS Fabr.
One specimen only, flying in the sunshine,@n March 2, 1906.
Does not occur on Christmas Island, but the genus is represented
there by JZ. dolosus Cdz. and M. litura Cdz.
HETEROMERA.
(11) Opatrum sp. (near simplex Fabr.).
A common insect, found mostly under the bark of the coconut
palms. Does not occur on Christmas Island, where the genus.
is represented by O. dubiwm Arrow.
(12) Ceropria inpuTA Wied.
Very abundant on all the islands. Lives in rotting wood.
The genus is not present on Christmas Island.
(13) SESsINIA sp.
Native name, “ Madu.
A very common insect. Found on all the islands, and freely
attracted to light. It is the subject of great variations in size.
Well known to the inhabitants, Malay and European, as pro-
ducing an acute dermatitis by contact, and an acute urethritis
when swallowed in drinks; for this reason drinks are always
carefully covered over to prevent the beetles from falling in.
Concerning these properties of the insect there is no doubt, and
although I have failed to produce the urethritis by swallowing a
decoction of two beetles in a glass of whiskey and soda, I have
met with several cases in which its development is definite, and
have frequently seen the cutaneous lesions.
This species would appear to be very similar to, if not identical
with, S. andrewst Arrow, from Christmas Island; and of that
insect Dr, Andrews says that it exudes an oily liquid, “ which is.
considered by the residents to have most injurious properties ”
(‘ Monograph of Christmas Island,’ p. 107).
LONGICORNIA.
(14) CerEstumM sIMPLEX Gyll.
Rare. Not found on Christmas Island, but two representatives
1909. ] FAUNA OF COCOS-KEELING ATOLL. 153
of the genus—(C. guadrimaculatum Gahan and C. nigrum Gahan—
were taken there by Dr. Andrews. >
(15) CaLocycius ANNULARIS Fabr.
Only one specimen taken, flying in the sunshine, on March Ist,
1906. Does not occur on Christmas Island.
(16) Ruorica HoNESTA Pasc.
(17) Ruopica Binotara Gahan.
Both unique specimens. The genus is not represented on
Christmas Island.
Seventeen species of beetles therefore occur on the Cocos-
Keeling group, and only one—a cosmopolitan species—is definitely
known to be also an inhabitant of Christmas Island, although
the coleopterous fauna of that island embraces ninety-five species.
RHYNCHOTA.
By W. L. Distant, F.E.S.
Suborder HETEROPTERA.
Fam. PENTATOMID2.
Subfam. CypDNINz.
(1) GEoromus PYGM&US.
thus pygmeus Dall. List Hem. i. p. 120 (1851).
Geotomus pygmeus Sign. Ann. Soc. Ent. Fr. (6) 11. p. 51, t. 111.
f. 160 (1883); Dist. Faun. B. I., Rhynch. i. p. 98, f. 49 (1902).
Hab. Widely distributed: recorded from Ceylon, Bombay,
Burma, Andaman Islands, and generally distributed throughout
the Malayan Archipelago; found in China and Japan, and
recorded from New Caledonia and Hawaii.
Subfam. PENTATOMIN2.
(2) NEZARA VIRIDULA. :
Cimex viridula Linn. Syst. Nat. ed. 10, p. 444 (1758).
For full synonymy ef. Dist. Biol. Centr-Am., Rhynch. 1. p. 78
(1880).
This species is distributed throughout the Palearctic, Nearctic,
and Ethiopian regions, and over a large portion of the Neotropical,
Oriental, and Australian regions.
Subfam. ASOPINa.
(3) OECHALIA CONSOCIALIS,
Pentalatoma consocialis Boisd. Voy. Astrol., Ent. u. p. 630,
Geena nC 8 35):
154 DR. F. WOOD JONES ON THE [ Feb. 16,
Oechalia consocialis Stil, Enum. Hem. 1. p. 59 (1870); Schont.
in Wytsm. Gen. Insect., Fase. 52, p. 75, t. v. f. 12 (1907).
This species is recorded from, and not uncommon in, Australia,
New Zealand, and Tasmania. It is a well-known species in
Queensland.
Fam. ARADID®.
(4) BRACHYRHYNCHUS MEMBRANACEUS.
Aradus membranaceus Fabr. Syst. Rhyng. p. 118 (1803).
Brachyrhynchus membranaceus Stal, Hem. Fabr. 1. p. 96 (1868);
Dist. Faun. B. I., Rhyne. ii. p. 160 (1904).
Found throughout British India, Malay Peninsula, and
Malayan Archipelago.
Fam. REDUVIID®.
Subfam. NABIDINA.
(5) NABIS CAPSIFORMIS.
Nabis capsiformis Germ. in Selb. Rev. Ent. v. p. 132 (1837);
Dist. Faun. B. I., Rhynch. i. p. 400, f. 256 (1904).
Distributed in the Nearctic, Palearctic, Ethiopian, Oriental,
and Australian Regions.
Fam. CApsiD#®.
(6) Lyeus sp. 2
Two specimens of a species of this widely distributed genus.
[None of these species can be described as at all abundant on
the atoll, and only Vezara viridula is commonly met with; the
others are, for the most part, very seldom found. The whole
of the order collected on Christmas Island by Dr. Andrews was
not worked over at the time of publication of his Monograph ;
but of the four species described by Mr. Kirby, there is not one
that is common to Christmas Island and the Cocos- Keeling atoll.
So far as I know, none of these species has earned a distinct
native name, but the domestic representative of the family,which
is common in native houses, is called “ kutu basuk” = stinking
insect.—F. W. J.]
Suborder HOMOPTERA.
Fam. FuLGORIDA.
Subfam. RiIcANnrInz#.
(1) Nocopina BOHEMANT.
Ricania bohemani Stal, Freg. Eugen. Resa, p. 280 (1858).
Nogodina bohemani Melich. Ann. Hofmus. Wien, xiii. p. 305,
t. xiv. f. 9 a@ (1898).
om
1909. ] FAUNA OF COCOS-KEELING ATOLL. 13)
Originally described from the Keeling Islands.
(This is not a common insect, and only a few examples were
taken, all from the leaves of the Hibiscus trees on the seaward
side of Pulu tikus.—F. W. J.}
NEUROPTERA.
Species determined by W. F. Kirey, F.L.S., F.E.S.
ODONATA.
(Native family name, “ Kachapong.”)
(1) PANTULA FLAVESCENS Fabr.
A very abundant insect ; it flies about all over the lagoon, and
is present on all the islands. In April and May of 1906 the
whole atoll swarmed with dragonflies, but for some time previous
to that it had been very rare to meet with a single specimen.
In the early months of 1905 these insects were entirely absent
from the atoll.
(2) TRAMEA ROSENBERGIL Brauer.
First seen on May 16th, 1906, and during the fallowing week
it became abundant; but for nearly a year previous to this it
had not been seen, and its numbers soon diminished afterwards.
(3) ANAx Gurratus Burm.
This species also came first to the atoll in May 1906, and then
only about a dozen examples were seen.
None of these species is resident on the atoll, and there is
no open fresh water for them to pass their early stages in. All
are wind-borne waifs, and do not belong properly to the fauna
of the islands, although they are at times so conspicuous a
feature of it.
Pantula flavescens appears to fly over the surface of the sea
from choice, and not to be, by any means, an involuntary
traveller; on the two occasions on which I have sailed from
Java to the atoll, I have seen—on calm days—many dragon-
flies hawking about above the water. It is not easy to see what
they are seeking, and yet they do not seem to be flying about
merely at the sport of the wind. In November of 1906, whilst
lying 20 miles to the south of the island of Sumbawa, these
dragonflies were seen every day, during ten days of calm, to be
flying in all directions over the surface of the sea, and as great
numbers of butterflies and other insects were also to be seen,
it is probable that they were in quest of food ; any question of
their being blown from the land was quite impossible. In 1907,
during a spell of quite remarkable calm, dragonflies were
seen on practically every day of a voyage between Sydney and
Singapore.
156 DR. F. WOOD JONES ON THE [ Feb. 16,
On the atoll itself, Zramea rosenbergii was often seen to
copulate, and apparently to deposit its eggs in the salt pools, but
none of its larvee could ever be found. Pantula flavescens and
Anazx guttatus ave both visitors to Christmas Island.
PLANIPENNTA.
(4) HemeErosius ? sp.
Native name, ‘ Lalar ijou,” = green fly.
This is a very abundant insect ; it occurs on all the islands, and
is certainly resident. When alive it is bright green, and its
eyes are remarkably brilliant. On account of its very offensive
smell when crushed, it is very well known. It is common at all
seasons of the year.
(5) Isoprera.
Native name “ Gegat.”
One species of ‘“ white ant” is very abundant in the wood-
work of dwelling houses. It is noteworthy that the natives
never include this insect under the title of Samut,—the family
name for ants.
ORTHOPTERA.
Species determined by W. F. Kirsy, F.L.S., F.E.S.
ForFIcuLID!.
(1) ANISOLABIS ANNULIPES Luc.
A common species found upon all the islands. It does not
occur on Christmas Island.
Buarripa.
(Native generic name, ‘‘ Kerklak.”)
(2) BLATELLA GERMANICA Linn.
(3) Auuacta NoctuLata Stal.
(4) Axxacta optusata Brinn.
(5) Lopoprera sp.
(6) Monyrria sp. (young larva).
(7) LevcopH@a suRINAMENSIS Linn.
All these species are common, and are mostly found beneath
the bark of trees, or in the dwelling-houses. Only Lewcophoa
surinamensis occurs on Christmas Island.
(8) PERIPLANETA AMERICANA Linn.
Common in store-houses ; introduced by ships.
1909. ] FAUNA OF COCOS-KEELING ATOLL. 157
ACHETID®.
(9) GRYLLODES SIGILLATUS Walk.
Native name, “ Orong Orong.”
Most commonly taken near to dwelling-houses; not very
abundant.
(10) OrNEBIUS sp.
A very abundant species. It lives in the bushes of ironwood
(Cordia subcordata), and passes its early stages in a rolied-up leaf.
Neither of these species occurs on Christmas Island.
(11) GryLLAcris sp. near signifera Stal.
Native name “ Chingkrek.”
This species also passes its early stages in the leaves of the
Cordia. When adult it appears to be carnivorous; it has
exceedingly powerful jaws, and is credited with waging war on
the large centipedes. When put in a box with a centipede it
certainly fights with great vigour, and though it bites the
centipede, the fights that I have arranged ended fatally for the
Gryllacris,—but I believe that this is by no means invariable.
This species does not occur on Christmas Island, but the allied
G. rufovaria takes its place. The antenne are 150 mm. long.
(12) Puisis pecrinata Guer.
A fairly common species, usually found in the Cordia bushes.
It is a bright green when living. It is not found on Christmas
Island, but P. listeri, that occurs there, is very nearly allied to it.
(13) ConocEPHALOIDES SOBRINUS Bol.
The male is buff-coloured, and the female green, during life.
The species is abundant, and commonly lives in bushes. It is
very musical. Does not occur on Christmas Island.
Locustip&,
(Native name, “ Blalang blalang.”)
(14) AcRyDIUM, sp. near japonicum, Sauss.
Very abundant on all the islands. Feeds on the fronds of the
coconut palm. It varies greatly in size when adult, and is the
subject of great seasonal changes of abundance and rarity.
Tt is not found on Christmas Island.
Although fourteen species of Orthoptera are found on the
Cocos-Keeling atoll, and twenty-three occur on Christmas Island,
there is only one species—Leucophea surinamensis—that is
common to both places. Of this species Dr. Andrews took only
a single specimen,
158 DR. F. WOOD JONES ON THE | Feb. 16,
ARACHNIDA anp MYRIAPODA.
By A. S. Hirst, F.Z.8.
The Arachnida collected by Dr. Wood Jones include several
species which are well-known to be distributed through human
agency. Of the few remaining forms, two spiders and one
myriapod seem to belong to undescribed species, but it is
probable that they will prove to be Malayan or Kast Indian forms
which have been introduced through trade with these regions
or have reached the islands by natural means.
SCORPIONS.
(1) Isomerrus mMacuLATUS De Geer.
Native name, “ Klajingking.”
Cosmopolitan.
SPIDERS.
(Native family name, ‘ Laba laba.”)
(2) SMERINGOPUS ELONGATUS Vinson.
Cosmopolitan.
(3) PuysocycLus GLogosus Taczaon Taczanowski.
Widely distributed in the tropical regions of the world.
(4) NepHina IMPERATRIX C. K.
An adult female example, which apparently belongs to this
species, was collected by Dr. Wood Jones. It differs from Koch’s
description of the species in having the abdomen marked ventrally
with two dark median patches, the anterior one being square
and separated from the smaller posterior patch by a whitish line.
The British Museum possesses examples of this form from
Buitenzorg, Java, and from Australia (Keyserling Coll.)
(5) Hereropopa VENATORIA Linn.
Cosmopolitan.
(6) Liycosa n. sp.
(7) BaAvra n. sp.
MILLIPEDES.
(Native name, ‘‘ Kaki ribu.”)
(8) TRIGONIULUS nh. sp.
(9) OnTHOMORPHA COARCTATA Saussure.
(10) There is also a large Centipede—native name ‘“ Alipan ”—
no specimen of which reached England.—F. W. J.
1909. | FAUNA OF COCOS-KEELING ATOLL.
CRUSTACHA.
(Native family name, “ Kapeting.”)
159
The collection of this order was not at all a representative one,
such specimens as were brought home have been determined by
Dr. W. T. Calman, F.Z:8.
BRACHYURA.
(1) CaRUPA LHVIUSCULA.
(2) LeproDIus SANGUINEUS.
(3) LroxaNrHo PUNCTATUS.
(4) LiomERA PUBESCENS.
(5) AcraA FOSSULATA ?
(6) CARPILIUS MACULATUS.
(7) HURUPPELLIA ANNULIPES.
(8) PsruDozIUS CAYSTRUS.
(9) ERipHiA LAVIMANA.
(10) MELIA TESSELLATA.
(11) GeELAsIMUS sp.
(12) CARDISOMA HIRTIPES.
(13) GECARCOIDEA LALANDEI.
(14) LroLopHus PLANISSIMUS.
(15) ScHIZOPHRYS ASPERA.
(16) CALAPPA HEPATICA.
Tn addition to this list there are several members of the genus
Ocypoda that live on the lagoon beaches, and several of the genus
Leptograpsus, of which one, the native name of which is ‘‘ Kapeting
traleg,” is eaten in the islands; a member of the genus Geograpsus
is one of the common land species.
ANOMURA.
(1) REMIPES TESTUDINARIUS.
(2) BirGus LATRO.
(3) C@NoBITA CLYPEATUS.
(4) Ca@NoBITA RUGOSUS.
(5) Ca@xoBitA PERLATUS.
(6) CALCINUS HERBSTII.
(7) CLIBANARIUS CORALLINUS.
160 MR. F, E. BEDDARD ON THE [ Feb. 16,
CARIDEA.,
(1) ALPHEUS STRENUUS.
STOMATOPODA.
(1) GONODACTYLUS CHIRAGRA.
VERMES.
There is one species of earthworm found in all the islands,
but unfortunately no specimen reached England.
+
2. Contributions to the Anatomy of certain Ungulata, in-
cluding Tapirus, Hyrax, and Antilocapra. By FRANK
BE. Bepparp, M.A., F.R.S., F.Z.8., Proseector to the
Society.
[Received January 28, 1909. ]
(Text-figures 10-19.)
T have during the past three or four years dissected and studied
various organs in a number of Ungulate animals, to which group
T have indeed paid special attention in view of the fact that for
obvious reasons these mostly large animals have been compara-
tively little examined. Indeed, of two or three of the species
of Antelopes to which I shall call attention in the following pages
there is, so far as I am aware, absolutely no knowledge of the
structure of certain of the soft parts. A good deal of work has
been done upon this group lately by Dr. Einar Loénnberg,
C.M.Z.S8., and upon some of the genera with which I occupy
myself in the following pages, viz., Cephalophus, Madoqua, and
Gazella. I am able, however, to add something to the large
number of facts which Dr. Léinnberg has accumulated in his
various papers to which reference will be made in the proper
places. The notes which I have to communicate to the Society
may be arranged under the following headings, viz. :—
(1) On the absence of a Pleural Cavity in the Indian Tapir,
SGIE
(2) Notes upon the Brain and some other points in the
Anatomy of Hyrax, p. 162.
(3) On the Existence of a new Skeleto-visceral Muscle in the
Pygmy Hog (Poreula salvania), p. 170.
(4) Notes upon the Anatomy of the Prongbuck (Antilocapra
americana), and on the Colic Helicine in some Artio-
dactyles, p. 172.
(5) Some Notes upon the Anatomy of Aladoqua phillipsi,
ay ltexsh
(6) The Brain of Babyrussa alfurus, p. 192.
(7) Résumé of new facts, p. 196.
OS ANATOMY OF CERTAIN UNGULATA. 161
(1) On the absence of a Pleural Cavity in the Indian Tapir.
In dissecting, some months since, an example of the Indian
Tapir (Vapirus indicus) which died in the Society’s Gardens, I was
struck by the close attachment of the lungs to the wall of the
chest-cavity. This attachment was by means of fine and multi-
tudinous strands of a glistening appearance precisely like a much
subdivided mesentery, and was very complete; so much so that
it would be quite fair to speak of the pleural cavity as practically
non-existent in that animal. It is unnecessary to dwell upon the
fact that the lungs were obviously not much moveable owing to
this firm connection with the walls of the cavity containing them.
Inasmuch as adhesions of a similar kind are by no means in-
frequent as a pathological phenomenon, it behoved me to decide
very carefully whether, in the example of the Indian Tapir which
exhibited the condition referred to, there was not some disease of
the lungs or the pleura, or of both, which would account for this
remarkable state of affairs. Although I have naturally accumu-
lated during my long tenure of the Prosectorship of this Society
some knowledge of morbid appearances and structures, I called in
my then colleague, Dr. Seligmann, to my assistance, who confirmed
my opinion. There can therefore be but little doubt that the con-
dition described above in Vapirus indicus is the normal condition.
It would appear, however, that this condition has been observed
in the same animal and has been put down to the effects of disease.
For Dr. Murie in his account of the anatomy of this Tapir *
wrote:—‘‘ The examination of the thoracic cavity exhibited the
residual effects of pleurisy, there being fluid exudation and
adhesions between the parietes and the lungs.” It is not so very
likely that so competent an observer as Dr. Murie would be
mistaken as to the presence of disease; but it must be borne in
mind that a normal attachment of the lungs to the parietes would
not prevent the lungs from being invaded by disease !
Again Dr. Cantor says7 that his specimen of Vapirus indicus
died from inflammation of the lungs, but he gives no detail. On
the other hand, neither Poelman = nor Sir Everard Home § say
anything bearing upon the present matter.
The interest of this remarkable obliteration, partial at any rate,
of the prediaphragmatic ccelom in Z'apirus indicus largely depends
upon the fact that the same state of affairs exists in the Elephant.
Quite recently Boas || and more recently still Giard 4] have shown,
in accordance with the statements of others (Miall & Greenwood,
&e., quoted by Giard), that the lungs are firmly adherent to the
* “On the Malayan Tapir, &c.,” J. Anat. Phys. vi. 1872, p. 139.
+ “Cat. of Mammals inhabiting the Malayan Peninsula and Islands,” Journ.
Asiat. Soc. Bengal, xv. 1846, p. 266.
{ Mém. Ac. Roy. Belg. xxvii. 1853. § Phil. Trans. 1821, p. 272.
|| “ Fehlen der Pleurahdhlen beim indischen Elefanten,’ Morph. Jahrb. Bd. xxxv
1906, p. 494.
@ “L’Eléphant de Afrique a-t-il une cavité pleurale,” C. R. Acad. Sci. t. exliv.
1907, p. 306.
Proc. Zoou. Soc.—1909, No, XI, il
162 MR, F. E, BEDDARD ON 'THE [ Feb, 16,
parietes by matted ligamentous cords in both Llephas africanus
and #. mavimeus. M. Giard goes on to suggest that the same may
be the case with /7yrax capensis, quoting “apparently confirmatory
evidence from the well known memoir of George upon that
animal, tL myself am unable to contirm this suggestion from my
dissections of several examples of both J/Zyrax capensis and
/, dorsalis, Still there remains the Elephant, which seems most
undoubtedly to resemble the Indian Tapir in this remarkable
specialisation, Lt is difficult to form an opinion as to whether
any stress is to be laid upon the fact that both these animals are
Ungulates.
(2) Votes upon the Brain and some other points in the
Anatomy of Hyrax.
The Society acquired some months ago the temporary posses-
sion of a considerable number of examples of LHyraxe capensis,
which were all about half-grown. ‘The measurement of one
individual (and all were of about the same size) was a foot or
so in length. The dissection of so large a series of specimens has
enabled me to add some facts to what is already known about the
structure of this primitive Ungulate. I she ull deal with sueh
organs as T have specially investigated seri@éim, and shall in some
cases be able to compare ‘them with the corr esponding organs of
iH. dorsalis, of which I have dissected two examples.
A Tygos ve in.—l have already deseribed * the condition of this
vein in Zyrax and quoted the descriptions of others. Tt is
evident from what has been written on the subject, that the
prevailing arrangement in /Zyra@v is the presence of a single
azygos only, and that on the right side. In all the examples to
which the present note refers, thisarrangement of the azygos was
found. The comparative constancy of the azygos, Upon ‘whieh I
have already ventured to insist, is thus established for the present
species.
The Brain of A yrax has been figured by several anatomists.
Thus Serres gives in his ‘ Anatomie ‘Comparée du Cerveau’ fF two
views of the brain. Later Dareste reported { upon three brains
which were not very well preserved. Of these he wrote that they
*seloignaient du type des Pachydermes pour se rapprocher de
celui des Carnivores "—no hint, here, it will be noted, of any
Rodent attinities. Some years later Gervais § figured a cast of the
brain in his well-known paper upon the mammalian brain, The
most complete series of figures known to me of the brain of this
*Subungulate ” are those of George in his monograph || wpon
Hyrex. The brain is figured in many views and very thoroughly.
The brain of Wyrax is also figured by Sir Richard Owen in his
* ©The Azygos Veins in Mammals,” P. ZS. 1907, p. 193.
it 1S2k 1827, pl. 15. figs. 269, 278.
t Ann. Sei. Nat. (4) i. 1855. § Journ. de Zool. 1872.
| * Monographie du Genre Daman * Ann. Sei. Nat. (6) i, 1874, pl. xvi.
1909. | ANATOMY OF CERTAIN UNGULATA, 163
text-book*, The most recent account known to me is that of
Dr. Elliot Smith in the Catalogue of the Physiological Series in
the Museum of the Royal College of Surgeons f.
{ have examined altogether fourteen brains of Tyrax capensis,
which permits me to contribute something to what is known of
the variations of the sulci in this Subungulate. None of these
brains, 1 should add, was among those studied by Dr. Elliot
Smith. This anatomist has studied seven brains and finds “ that,
the features of the brain vary greatly in different specimens.”
In addition to the brains of Myrax capensis, 1 have examined
one brain of /, dorsalis. J shall now proceed to review such
variations as are exhibited in the fourteen brains of JI. capensis,
commencing with the dorsal surface,
The splenial sulcus is figured by Elliot Smith as invisible pos-
teriorly upon the dorsal surface; its position for the last 5 mm.
or so of its length is mesial and lateral, and it is therefore not
to be seen when the brain is regarded from the dorsal surface.
This was also the case with four of the brains examined by
myself. In the others the sulcus could be seen dorsally for its
entire length. This was also very markedly the case with the
one brain of /T. dorsalis.
As a rule, the splenial sulcus is quite independent of the
coronary and lateral sulci. The blood-vessel running along it,
however, is formed by two branches, one behind the other, to the
vessel running along the coronary sulcus. ‘These naturally
dahil indent the surface of the brain. But in two specimens
only out of the entire series, and in them only on the right side,
was there a distinct furrow joining the splenial to the posterior
end of the coronary. his furrow represe nts an extension of the
small notch figured by Elliot Smith? as lying between the
coronary and lateral sulci. The aicngianel completion of this
fissure is of course suggestive of other Ungulates §. he coronary
fissure does not always stop short of the transverse fissure or notch
in the middle of the length of the brain which has just been
referred to. This was, however, the case with six brains. In the
others the coronary fissure was connected by the transverse fissure
with the suprasylvian.
In the single brain of Myrax dorsalis there was no connection
between the coronary and the suprasylvian.
The lateral fissure showed no variations except very slight ones
of length.
The lateral view of the brain is figured by Elliot Smith in two
examples, which differ very considerably. J have also found a
certain number of variations in the sulci of this region of the
brain, which are the following :—
In eight examples I found the suprasylvian fissure to be very
* © Anatomy of Vertebrates,’ vol. iil. fig. 96.
+ 2nd ed. vol. ii. p. 297. Dr. Elliot Smith quotes also a memoir by Turner
(J. Anat. Phys. xxv. 1891) and Krueg’s well-known paper; see also Elliot Smith,
Trans. Linn. Soc., Zool. (2) viii. p. 400.
{ Loe. cit. p. 297, fig. 168. § See below, p. 173, and text-fig, 11.
1%
164 MR. F. E. BEDDARD ON THE [ Feb. 16,
slightly bowed and not nearly reaching the rhinal fissure; and
the single brain of Hyrax dorsalis agreed with these. In the
remaining brains the suprasylvian fissure was carried on to the
rhinal. I found an orbital fissure in all of the brains, and in five
or six there were more or less well-developed traces of a preorbital
fissure which did or did not join the rhinal fissure and rarely
jomed the coronary above ; in the latter event the fissure did not
communicate with the rhinal.
On the temporo-occipital region of the brain, 7. e. behind and
below the suprasylvian fissure, T find the existence in var ying pro-
portionate degrees of three more or less parallel fissures which are
vertical in direetion—emd not of two only as figured by Elliot
Smith. The most anterior of these is, however, obviously repre-
sented by the lowermost section of the supr asylvian fissure figured
in one of Dr. Elliot Smith’s illustrations*. For I have found in
some brains where the suprasylvian fissure is as strongly bowed as
in the illustration referred to, that the suprasylvian fissure ends
anteriorly in a Y-shaped bifurcation of which in other brains one
or other limb may be entirely or partly absent. In the brain
figured by Dr. Elliot Smith 7, as in a brain among those examined
by myself, the upper anterior limb of the bifurcation is absent ;
whereas in the second brain figured by Elliot Smith it is, as I
also have seen, the lower part of the fork which las vanished.
These three fissures seem to me to vary from individual to
individual and on both sides of the brain of the same individual
very often. The principal variations in the specimens examined
by myself were the following :—The most anterior of the three
fissures was quite absent or represented by a small rudiment which
did or did not join either the rhinal or the suprasylvian fissure,
or finally it effected a complete junction between these two fissures.
The middle fissure which Elliot Smith letters in his drawings as
the Sylvian is sometimes longer, sometimes shorter; sometimes it
starts from the rhinal fissure and is sometimes unconnected with
that fissure. It is occasionally broken into two smaller fissures.
lying one above the other. More rarely, it is connected by a
transverse fissure (¢. ¢. one running horizontally) with the third
of the three fissures referred to and which Elliot Smith terms
postsylvian. This fissure again varies very much in length and
sometimes joins the suprasylvian. Very rarely it joins the rhinal
below.
These variations are shown in the accompanying figure (text-
fig. 10), where not only the differences between one brain and
another are exhibited, but also those between the two sides of the
same brain. J imagine that the series selected almost exhausts
the variations which can occur in this region of the brain. It will
have been observed in reading the foregoing remarks that nearly
all the sulci of the brain of Hyrax are subject to variation—
that indeed only the lateral furrow remains constant.
* Toe. cit. fig. 1771, p. 299. + Loc. cit. fig. 169, p. 298.
1909. | ANATOMY OF CERTAIN UNGULATA. 165
Arteries of the base of the Brain.—\ figured in the ‘ Proceed-
ings’ of this Society some years since* the chief arteries which
form or are derived from the circle of Willis in Hyrax capensis.
A number of the brains upon which I have just reported show the
Text-fig. 10.
Tateral view of the Sylvian region of eleven brains of Hyrax capensis.
Figs. 1, 2, 3, &c. represent the left side. Figs. la, 2a, 3a, &e. the right side
Each figure is bounded by the suprasylvian fissure above and the
rhinal fissure below.
cerebral arteries and enable me to amend in some particulars ana
to confirm in others the figure and description referred to. I have
there figured no communication between the basilar artery and
* P.Z.S. 1904. vol. i. p. 187, text-fig. 18.
166 MR. F. E, BEDDARD ON THE [ Feb. 16,.
the right side of the circle of Willis. It is clear, however, that
this specimen was so far abnormal; for in nine others (the only
ones in which the arteries were visible) both divisions of the
basilar artery were plainly present, thus completing the symmetry
of the circle of Willis.
I find that asa general rule the basilar artery is undivided upon
the medulla, as I have figured it in Hyrax *. But in two out of
the nine specimens the artery formed a loop, such as I have
figured in Zamandua, but longer. Such a bifurcation and re-
junction of the basilar artery is very common among Ungulates.
In each case the vertebral artery on either side joins the anterior
rhinal artery behind the loop. These arteries mark the end of the
medulla oblongata. Imay remark that a misinterpretation of the
woodcut illustrating the arteries of the base of the bra in
Equus in Messrs. Chauveau and Arloing’s treatise, led me _ to-
compare Hyrax especially with Hquus in the posteriorly situated
communication of the carotid with the circle of Willis. They are
as a matter of fact anteriorly placed in the Horse as in the
Rhinoceros 7, and posteriorly placed in Hyrax. In five examples
of Hyrax | found that the carotid became continuous with the
circle of Willis just opposite to the origin of the posterior cerebral
artery. The other specimen examined by me did not show the
relations of the artery with great clearness. I mention this matter
particularly since it is not dealt with in my original figure of the
arterial system of the brain. I find that the arteries at the base
of the brain in Hyrax differ in another particular, unless indeed I
have made an error in my former description. For I find that in
several brains the anterior cerebellar artery arises on each side
from the bifureate basilar artery, and not from the latter posteriorly
to its bifurcation. In spite, however, of these few additions to
the description of the arrangement of the arteries forming the
circle of Willis, it remains clear that Hyrax comes nearer in
respect of these structural peculiarities to the Perissodactyle than
to the Artiodactyle section of the Ungulates, a position upon
which I ventured to insist in the paper already quoted.
Cecum and gut of Hyrax dorsalis.—Of this species of Hyrax I
have recently dissected two examples which permit me to compare
certain organs with the corresponding organs in Hyrax capensis
and Hyrax syriacus. In the general subdivisions of the cecum of
H. dorsalis 1 find no cuatrences from those of H. capensis. When
the cecum is viewed on the opposite side to that on which the
ileum enters it, two muscular bands as in H. capensis are seen to
traverse the cecum, forming by their contraction two parallel
furrows upon the surface of the cecum; these ultimately join to
form a single muscular band and furrow. The junction is, how-
ever, considerably further forward in Hyrax dorsalis, at a point
in fact corresponding to the end of the first two-thirds of the
entire length of the cecum. Furthermore, while in H. capensis
* Loc. cit. p. 189, text-fig. 19 b.
+ Beddard & Treves, Prans. Zool. Soe. vol. xii. pl. xxxvil.
1909. | ANATOMY OF CERTAIN UNGULATA. 167
the czeco-colic ligament extends down to the end of the cecum
more or less along the inner of the two muscular bands only, this
ligament is much less extensive in H. dorsalis. It lies, as in
Hf. capensis, upon the outer of the two muscular bands of the
cecum, 7. e. that on the colic side of the cecum, but does not
extend back even so far as the point where the two bands unite to
form a single muscular band.
The cecum of H. dorsalis then, like that of H. capensis, shows
on the ventral side two longitudinal teeniz where only one perhaps
would have been expected. I take this to mean that the space
included between these two muscular bands is the ventral surface
of the median chamber of the cecum, which is so apparent
dorsally between the two lateral chambers and is externally
divided from them by two muscular bands bearing blood-vessels
which are continuous in front with the ileo-colic mesentery.
When the cecum is dilated with fluid, these bands are seen to
produce much puckering of both the median and lateral portions
of the cecum. When the cecum is cut open these three chambers
are recognisable, but there are no permanent folds dividing them ;
the walls of the cecum can be pressed flat. On the other hand,
there are two anterior chambers of the cecum, right and left,
which are divided from the rest of the cecum by permanent
though low folds which cannot be stretched out flat; they are
actually valve-like thickenings of the walls of the cecum. These
lie in front of the entrance of the ileum and closely embrace the
dilated commencement of the colon, the bands bearing blood-
vessels already referred to upon the dorsal side of the posterior
region of the czecum divide them off from the median “ chamber”
of the posterior region of the cecum.
I have recapitulated these facts, firstly to show the general
resemblance of H. dorsalis to H. capensis in the form and sub-
divisions of the cecum, and secondly to emphasise another
difference between the two species. This is the extension back-
wards of the median dorsal chamber beyond the two lateral
chambers, so that it can be seen on the ventral aspect of the
cecum in Hf. capensis. This is not the case with H. dorsalis.
There are thus two quite distinct differences between the ceca of
these two species. The two chambers already referred to which
embrace the terminal section of the cecum or the beginning of
the colon, according to the interpretation preferred, are spoken of
by Lonsky * as the “ mittlere Etage” of the cecum, lying in fact
between the lowest story which forms the blind end of the cecum
and the upper story out of which emerges the colon. Jam inclined
to consider that this region is potentially double—a meeting of
the actual folds which delimit each half would convert each half
into a tube—as indeed it appears to be externally. In this case
a comparison may be fairly made with the two cecal appendages
of the Manatee’. Lonsky has indeed compared the cecum of
* Jen. Zeitschr. xxxvii. 1903, p. 580.
+ Maurie, Trans. Z. S. viii. pl. 23, fig. 28; & Beddard, P. Z.S. 1897, p. 50, fig. 3.
168 MR. F. E. BEDDARD ON THE [ Feb. 16,
Manatus with that of Hyrax, but not in regard to the paired
outgrowths. He has rather emphasised the median chamber in
which they meet, and has compared this with the two chambers of
the cecum of Hyrax as I consider them together with the median
space common to the two. Lonsky observes that the ileum enters
the middle chamber of the ceecum and that it is without an ileo-
cecal valve. ‘The latter statement is certainly not true of H. dor-
salis, the valve was quite obvious in the two specimens which I
have dissected. Furthermore it appeared to me that the ileo-
cecal orifice lay in this species of Hyrax in the lower and largest
chamber of the cecum. The raised fold which separates off from
this latter subdivision the two diverticula (as I regard them) can be
distinctly seen to join the posterior lip of the ileo-czecal valve, @. é..
that nearest to the colon. On the other hand, there is undoubtedly
on each side a branch of this fold which passes to the other side
of the ileo-cecal orifice. The posterior fold, however, appeared
to me to be the most important. The question is obviously a
difficult one to be decided. Finally (as regards the alimentary
tract) I may remark that the present species, Hyrax dorsalis,
differs from H. capensis and agrees with H. syriacus as described
by Lonsky, in possessing a longish cecal appendage of the colon
lying between the true cecum and the paired colic appendages.
Nearly up to the very tip of this diverticulum, which is about an
inch long, is attached the ceco-colic ligament, the other attach-
ment of which to the cecum has been already described.
Although Hyrax capensis does not possess this diverticulum, there
is certainly a slight dilatation of the colon where the czco-colic¢
ligament is fixed. It does not appear to exist in Hyrax brucei*.
The Testes of Hyvax dorsalis and H. capensis.—In all the speci-
mens of both of these species which I have examined, the testes
(which are well known to be invariably intra- -abdominal) are partly
enveloped by a freely hanging membrane of considerable cireum-
ference, of which I can find no description in recent memoirs
dealing with this Ungulate, and which therefore is at least not
well known. The most recent and most elaborate survey of the
genitalia of Zyrax (both male and female) known to me is in the
memoir of Lonsky already referred to T
Neither in the text nor in the figures illustrating it { does
Lonsky refer to the membrane to which I shall presently refer
at greater length. This structure has been described and figured
by Pallas §, whose description runs as follows :—‘‘ Testes... . et
margine libero coronati membranula, seu ala e peritonzeo facta et
adipe striata, quee vasa a spermaticis accipit.” It is illustvated by
a figure || which does not, however, represent quite accurately the
conditions which are observable. The sperm-duct is represented
* Chapman, P. Acad. Philad. 1904, p. 476:
+ Jen. Zeitschr. xxxvii. 1903, p. 612 &c.
{ Loe. cit. pl. 29. figs. 4, 5, 7.
§ Spicilegia Zoologica, Berolini, 1767, Fasc. 2. p. 29. || Tab. ii. fig. 11.
1909. ] ANATOMY OF CERTAIN UNGULATA. 169
as expanding into the freely floating membrane, and thus sug-
gesting unduly the funnel of an oviduct. This almost inclines
one to the belief that Pallas had confused the epididymis with
the membrane in question, though the phrase quoted above seems
to negative this belief. It seems, however, that later writers
have not referred to this structure, for the most part at any rate.
The classical memoir of Brandt * contains no account of the
membranous appendage of the testes, nor is it figured. Indeed,
this anatomist has only figured the female organs. Nor is it
represented by George t or Chapman. In both Hyrax dorsalis,
and H, capensis there is invariably (so far as my experience goes)
a loose fold of membrane, depending freely into the body-cavity,
of a reddish colour in parts at least, and with an irregular
edge, which is attached to each testis. This varies in size in
different individuals (at any rate, it varied in two examples of
Hyrax dorsalis) and is restricted to a less, or expands to be
attached toa greater, area of the testes. There are also differences
on the two sides of the body.
One of the most recent memoirs dealing with the anatomy of
the male organs of the Elephant is that by v. Mojsisovics upon
the African Elephant $. He describes ‘‘ Ein nahezu vollstindiger
Peritonealiiberzug heftet sie [7. e. the testis] an die mediale Seite
der Niere etwas unterhalb des Hilus. Cuvier vergleicht dieses
Aufhingsband der Form nach mit emem Lig. uteri laterale.”
I do not, however, think that this is anything more than the
ligament to which I have referred and which unites testis and
kidney in Hyrax. The late Dr. Morrison Watson || says nothing
of any such free floating membrane as that which characterises
Hyrax in the Indian Elephant. But Camper 4 speaks of “ La
membrane qui les enveloppe formoit des deux cotés plusieurs
franges garnies de longues appendices en forme de petits epiploons.”
The figure ** illustrating this appears to me to be of a structure
comparable to the testicular membrane which I here describe in
Hyracx.
These variations suggest an organ which is no longer functional ;
and it is in any case difficult to see what can be the actual use in
the economy of this floating sheet of membrane. It seems to be
quite distinct from the narrow and tightly stretched ligament
which joins the testis to the anteriorly lying kidney. It is, in
fact, associated with the testis, and not with the kidney. The
close association, its reddish colour, and fimbriated edges suggest
very much the appearance of and association with the ovary of
the oviducal funnel in Marsupials. And it is possible that this
structure is really the equivalent of an oviducal funnel persist-
ing in the male. There is, however, another possibility. The
* Mem. Ac. St. Peterb. (7) xiv. 1869.
+ Ann. Sci. Nat. (6) 1. 1874. t P. Acad. Philad. 1904, p. 476.
§ Arch. f. Naturg. 1879, p. 82. || J. Anat. Phys. vii. 1873, p. 60.
|
€ Description anatomique d’un Elephant male. Paris, 1802, p. 55.
*k. Loc. cit. pl. v. fig. 1.
170 MR. F, E, BEDDARD ON THE [ Feb. 16,
membrane in question may correspond to a gubernaculum testis
diffusely developed. This, however, appears to be less likely.
By many zoologists the Hyrax is regarded as forming with the
Hlephant a group, Subungulata, lying at the base of the Ungulate
series. Recently opinion appears to be veering away from this
view of the affinities of the animal. Lonsky, already quoted, sees
in the alimentary tract and genitalia “the most resemblance” to
the Rodents; while Erich Fischer * holds that the carpus and
tarsus are not derivable from a primitive Ungulate such as
Phenacodus, but since they combine the characters of Rodents
and Ungulates are perhaps more to be compared to those of the
Tillodontia or primitive Toxodonts. Finally, Assheton f cannot
see in the placenta any likeness to that of any Ungulate or
Hlephant, but compares it on account of its “‘ cumulate” characters
to that of the Insectivora, or even to that of the Primates. It is,
therefore, not without interest to note that in the characters of
the testis Hyrax shows a likeness to one of the few other Eutherian
Mammals in which the testes are completely abdominal in position
throughout life, viz., Hlephas. Lonsky denies any special resem-
blance in form, but says nothing of the testicular membrane with
which I am at present concerned.
(3) On the Existence of a new Skeleto-visceral Muscle in the
Pygmy Hog, Porcula salvania.
Among the Mammalia the only skeleto-visceral muscles, 7. e.
muscles which pass from some part of the bony skeleton to some
viscus or viscera, are confined, so far as I am aware, to the
diaphragm and to muscles in the hyoid region. Nor are such
muscles very common among the lower vertebrata. We have the
slips of muscle which run from the ribs to the surface of the lung,
and the two muscles connecting the windpipe with the skeleton,
in Birds; there are certain muscles connecting the vertebral
column and the cesophagus and lungs in Frogs; the ventral sheet
of muscle reaching the stomach and liver in Crocodiles and the
Lizards of the genus Varanus ; muscles in the mesentery of various
Saurians, and the tongue-muscles in all of these animals. These
are at least the principal instances of such skeleto-visceral muscles
among Reptilia, Amphibia, Crocodilia, and Aves.
It is, therefore, interesting and not without importance to be
able to add to this series a quite new muscle of the same kind in
the Pygmy Hog, Sus, or Porcula, salvania. The anatomy of this pig
was carefully described by Dr. J. G. Garson some years since in
the ‘ Proceedings’ of this Society t. TI cannot find in that account,
which is quite comprehensive as regards the viscera, any mention
* Jen. Zeitschr. Bd. xxxvii. 1903, p. 691.
+ Phil. Trans. B, vol. 198, 1906.
{ P. Z. S. 1883, p. 413. This, moreover, is described as the “first part.” The
second part, dealing with muscles, has not I believe appeared.
19093} ANATOMY OF CERTAIN UNGULATA. 171
of a peculiar muscle arising from the vertebral column in the
middle line and running to be inserted on to the cesophagus.
The direction of the muscle is almost transverse, and it arises
from the centre of either or both of the seventh and eighth dorsal
vertebre. It les opposite to the second branch of the azygos
vein, which in this animal is developed only upon the left side.
A nearer examination of this muscle where it ends upon the
cesophagus shows that it is continuous with a distinct and easily
separable muscular sheath of the esophagus. The muscle is so
large that it could hardly escape being seen if the thorax of the
species were examined by anyone. I 1 imagine, therefore, that the
reason why it is not described by Dr. Garson is that he ‘intended
to describe it later in the second part of his memoir dealing with
the bones, the muscles, and nerves of Porcula salvania. After
finding this peculiar and very obvious muscle in the Pygmy Hog,
I looked for it in some other Suide, but so far with negative
results. I did not find the vertebro-cesophageal muscle in
Dicotyles tajacu, or in Phacocherus cthiopicus. It was also
apparently absent in a young Sws cristatus, which completes the
list of Suidee which I have examined up to the present date for
this particular structure. Nor has a superficial examination of
other mammals shown anything of the kind to be present. I am
at least convinced that no such large muscle in the same situation
or hard by exists in a large number of mammals. Unfortunately
I neglected to note the minute structure of this muscle. I pre-
sume, however, from its large size that 1t consists of striated fibres.
I have described in the heading to this section of the present
communication to the Society the muscle just described as ‘ new.”
I believe it to be so. But at the same time something of the
kind has long been known to exist at any rate in man.
Thus Prof. Birmingham writes * :—‘ The longitudinal fibres
[z. e. of the cesophagus] are often joined by slips of unstriped
muscle, or elastic fibres, which spring from various sources,
including the left pleura Consaep Cunningham), the bronchi,
back of trachea, pericardium, aorta, &c. These slips assist in
fixing the cesophagus to the surrounding structures in its passage
through the thorax, and have been aptly compared to the tendrils
of a climbing plant.” It is plain that these shps of muscular
tissue also correspond to similar slips often very numerous and
closely adjacent which occur in the corresponding and in other
situations in the mesenteries of Saurians. A massing of these
would produce a muscle like that which I have just described in
Porculay salvania. This subject, however, evidently requires
further study. All that I can say at present is that there are
certainly some mammals in which a large vertebro-cesophageal
muscle like that of Porcula salvania does not exist.
* Textbook of Anatomy; edited by D. J. Cunningham. London & Edinburgh,
1902, p. 991.
+ It is true that Dr. Garson saw no reason in view of the structure of the animal
for retaining the generic name. I have, however, described a new character which
may go towards justifying its retention.
172 MR. F. E. BEDDARD ON THE | Feb. 16,
The sheath of muscles derived from this vertebral slip in
Porcula salvania continues on to, or to the neighbourhood of, the
diaphragm. As already mentioned, I neglected to ascertain.
whether this sheath and the muscular slip from which it is
derived were composed of striated fibres. Supposing that they
are—the most likely supposition, as I think—it may be of interest
to compare this sheath with that of other mammals. In a con-
siderable series of papers the late Mr. Gulliver * recorded a striate
sheath to the esophagus which in man may be regarded merely
as a slight extension backwards of the striated fibres of the
pharynx, but which is a more important matter in, for example,
Rodents and Ungulates. In them the striated layer continues
back to the hinder end of the esophagus. It is possible that this
state of affairs is traceable to the attachment of a muscle like that
described in Porcula, the greater part of which has disappeared
in other types, leaving only the cesophageal portion.
(4) Notes upon the Anatomy of the Prongbuck (Antilocapra ameri-
cana), and on the Colic Helicine in some Artiodactyles.
In the most recent paper known to me upon the American
Prongbuck Antelope +, Mr. Lyon enumerates the very abundant
sources of information respecting the horns of that anomalous
Antelope, and in discussing the systematic position of Antilocapra
refers to Dr. Murie’s account of its anatomy in the ‘ Proceedings’
of this Society $. I believe that this latter paper contains all that
is known up to the present time respecting the visceral structures
of the animal with which I am concerned in the present com-
munication to the Society. There is thus much less consensus of
opinion upon the general anatomy of this Bovine than upon the
nature of the horns, concerning which Mr. Lyon refers to no less
than 19 separate memoirs or references in larger works. I
endeavour in the following pages to supplement Dr. Murie’s paper
by some notes upon certain organs which he has not described or
not fully described.
The Azygos veins of Antilocapra are on the plan of those of the
typical Artiodactyla, meaning in this case, by “ typical,” the
majority. On the left side is the principal vein. This is formed
of an azygos which meets a precardinal vein running from before
backwards to meet the azygos or postcardinal. The former is not
long. The latter is as usual rather long and extends a moderately:
long way back towards the diaphragm. The point of junction of
the two veins and of their connection with the heart is opposite
to the fifth rib (there are altogether thirteen pairs of ribs). As.
is the case with other Artiodactyles, as a rule there are also veins
on the right side belonging to the azygos series, and of these there
are two which open, the one behind the other, into the precaval
* P.Z.S. 1869, p. 249, and, P. Z. S. 1870, p..283, with other papers quoted.
_*+ “Remarks on the Horns and on the Systematic Position of the American
Antelope,’ P. U.S. Nat. Mus. xxxiv. 1908, p. 393. t P. ZS. 1870, p. 334.
1909. | ANATOMY OF CERTAIN UNGULATA. 173
vein. The first of these, which is the superior vertebral vein,
supplies the first intercostal space; the next vessel is formed by
the union of two twigs which supply the next two intercostal
spaces. It will be seen that in detail the azygos system of this
Antelope differs from that of other forms. But it is at present
perhaps premature to compare the genus in this respect with
other Cavicorn or Cervine ruminants.
The Brain.—I believe that the brain of Antilocapra has not
been described, though a “ cast of the cranium” is referred to in
Text-fig. 11.
Brain of adult Prongbuck (Antilocapra americana), dorsal view. Nat. size.
a. Coronal sulcus. d. Suprasylvian. ¢. Lateral sulcus. £ Hntolateral sulcus.
the Catalogue of the Museum of the Royal College of Surgeons*. I
describe in the present communication a brain of an adult Anf#ilo-
capra americana recently living in the Society’s Gardens. The
outline of this brain as seen from the dorsal side is remarkable
for the squareness of the anterior portion in front of the fissure
of Sylvius, as seen in the accompanying figure (text-fig. 11) and
as referred to under the description of the brain of JMadoqua
on p. 190.
The splenial sulcus is not apparent upon the dorsal surface of
the brain ; its relations with the crucial sulcus anteriorly seem to
me to be remarkable and unlike those of some other Artiodactyles.
an Ve, BB);
174 MR. F. E. BEDDARD ON THE [| Feb. 16,
For example, in the brain of Madoqua phillipsi, which will be
shortly described in the present communication, the splenial
suleus is continuous anteriorly with the crucial suleus and does
not communicate with the coronal sulcus, which sends a branch
osteriorly to the middle line. Precisely the same arrangement
occurs in the Sheep, as figured by Elhot Smith. On the other
hand, in the Pig tribe, as is well known, and as will be also
pointed out in the present communication to the Society, the
splenial fissure 1s continued into the coronal. In the Babyrussa
there would seem to be a continuation of the splenial imto both
these fissures. The crucial fissure and the fissure lying behind it,
which represents the inner and posterior limb of the sulcus coro-
nalis, are as plainly shown in Antilocapra as in Madoqua (cf. text-
figs. 11 & 16) and in other genera *. The crucial fissure on one side
the right) joins the coronal, and on the other though continuous
with the coronal it does not reach the middle line of the brain.
The coronal behind is also asymmetrical, but in a different way ;
on the right the fissure is separated by a bridging convolution
from the junction of the coronal and the branch of the supra-
sylvian. On the left there is complete continuity.
Tt is this coronal fissure which is continuous with the splenial,
and not the crucial as in Ovis. The lateral fissure proper, which
T take to be the middle one of three lateral fissures, is remarkable
for the fact that at the end of its first third (about) there is a
strong branch at right angles on the outside, and also some indi-
cations of a similar branch at the same spot on the inside, the two
together forming a cross branch. The ecto- and entolateral sulei
run forwards to nearly as far as this point. The entolateral
fissure, it may be observed, does not nearly reach the middle line.
The suprasylvian fissure is absolutely continuous with the coronal,
as in Cervus. It bifurcates posteriorly in the occipital region
intoaY. There are two marked descending furrows, one in front
of and one behind the Sylvian. The genwal fissure is not con-
tinuous with the splenial (intercalary).
In addition to the brain of the adult Prongbuck just described,
IT have had the opportunity of studying two fetal brains
extracted from the mother, but of nearly if not quite full-time.
Unfortunately they were both rather damaged at the time, but
T have been able satisfactorily to compare certain of the sulei with
those of the large brain. The strong branch of the lateral sulcus
running at right angles to it, which seems to me to be so charac-
teristic of Antilocapra, is quite obvious in both of these brains
and on both sides of the brains. I have mentioned particularly
the fact that the entolateral suleus does not reach the middle line
of the brain. In one of the feetal brains the relationship was the
same; inthe other, however, the entolateral sulcus of the left side
only did dip over the mesial edge of its hemisphere. As to the
continuity of the coronal with the splenial sulcus I can only
* BE. g. Cervus, Elliot Smith, loc. cit. p. 329, fig. 190.
+ Holl, Arch. f. Anat. Phys. pl. xviii. fig. 10, & Elliot Smith, Zoc. ci¢. fig. 191, p. 329.
1909.] ANATOMY OF CERTAIN UNGULATA. 175
speak of one of the fetal brains, and in that there certainly was
this continuity. Furthermore, on one side of the same brain
(the right—I cannot say anything of the left) the crucial sulcus
was continuous with the coronal.
Muscular Anatomy.—\ have dissected some of the muscles of
one of the foetal specimens of Antilocapra, and have compared my
dissections with the comprehensive account of Ungulate myology
of Messrs. Windle & Parsons *. The latissimus dorsi cannot be
said to be a feeble muscle in this Ungulate. Its origins are, as
usual, in common with the trapezius and behind and contiguous
to that muscle from the spines of some dorsal vertebre, from the
lumbar fascia covering over the longissimus dorsi and ilio-lumbaris
muscles, and from three ribs nearer to their ventral than to their
dorsal ends—from the sides of the thorax in fact. A little way
down the scapula the muscle diverges from the trapezius and
passes under the broad triangular scapular head of the triceps to
reach the humerus. Just at this region it gives off a feebly
developed dorso-epitrochlear branch to the elbow, and is connected
with the teres major, in common with which it is inserted. Here
also the panniculus carnosus is attached to the latissimus.
The rhomboidei are, as is usual in Mammals, two in number
and composed of a more superficial and a deeper-lying muscle.
The superficial is the more anterior of the two, but only covers
the deeper-lying part of the muscle anteriorly. Its fibres are also
more antero-posterior in direction than those of the other. The
insertion to the edge of the cartilaginous supra-scapula commences
in contact with and posteriorly to the supraspinatus anteriorly,
and posteriorly ends a quarter of an inch or so in front of the
posterior commencement of the insertion of the serratus. These
two muscles, therefore, form a nearly complete semicircle (or
rather, semiellipse) of insertion on to the supra-scapula.
The serratus, as already mentioned, is quite contiguous ante-
riorly with the superficial rhomboid and arises from eight ribs,
the latissimus dorsi immediately following from the three ribs
which lie behind those eight. Its insertion on to the scapula is
both in front of and behind that of the rhomboidei.
The clavicular portion of the deltoid forms part of the great
Ungulate cephalo-humeral muscle. The spinous portion arises,
as in other Ungulates, not only from the spine of the scapula, but
also from the fascia covering the infraspinatus.
The subscapularis is fairly extensive. It arises from and covers
the greater part of the ventral surface of the scapula. There is
a very small gap left between it,and the semicircular insertion
of the serratus and rhomboidei. Anteriorly, however, a portion
of the subscapularis is somewhat distinct from the rest of the
muscle and leaves a tract of bone, not long, between its origin and
the insertion of the rhomboideus.
The supraspinatus extends over on to the ventral surface of
* P, Z.S. 1901, vol. ii. p. 626, and P. Z. S. 1903, vol. ii. p. 261,
176 MR. F, E. BEDDARD ON THE [| Feb. 16,
the scapula, where it comes into contact with the anterior edge of
the subscapularis muscle. It arches over the tendon of origin of
the biceps and is muscular at its insertion, with homer er a
glistening tendinous surface.
The infr aspinatus is distinctly divided at and for some little
way in front of its (therefore) double insertion into two layers, of
which one overlies the other.
Teres minor is a slender muscle ending in a long flat tendon
below the insertion of the spinous part of the deltoid, and there-
fore considerably below the insertions of the infraspinatus, which
are above that of the section of the deltoid referred to.
The teres major is a very much larger muscle, flat and strap-
shaped, and closely connected with the latissimus dorsi near to
insertion.
Biceps.—This muscle is single-headed, as in other Ungulates.
The strong tendon of origin seems to me to arise very distinctly
from the coracoid process, but also doubtless from the adjacent
margin of the glenoid cavity. Low down on the arm and at the
end of the insertion of the pectoralis, a stout fibrous band is given
off which runs diagonally towards the elbow. This is chiefly
connected with the lower end of the pectoralis, but is adherent to
the biceps where it crosses it. A little further down and just
before its insertion the biceps is connected with a stout tendinous
sheet running down the forearm, in the centre of which is a
stouter tendon which can be seen to end on the surface of the
biceps.
The brachialis anticus winds round the humerus in the way
that has been described in other Ungulates. Its insertion is.
remarkable for the fact that it is adherent by a tendon to the
stout tendinous plate running down the forearm, which has just
been described, in connection with the biceps before its definite
insertion on to the ulna.
Triceps.—This muscle consists of at least four very distinct
parts, unless it were better to term one of these anconeus. Of
these heads the scapular was precisely as described by Windle &
Parsons, arising from a large portion of the axillary border of the
scapula. When cut across transversely, there were considerable
indications of the division of this muscle into two layers, as also
noted by those authors. The imner humeral head seems to be
peculiar in some respects. It arises below the coracobrachialis
and immediately divides into two muscles, of which the posterior
is more or less cylindrical in form in the middle and circular in
transverse section, and is totally free from any connection with
the shaft of the humerus. The larger portion, that lying anterior,
has an origin from the humerus for about halfway down the
shaft of that bone. When it becomes free it is rejoined by the
other slip of the muscle just before mentioned, and the two form
a flat tendon at the internal condyle. The external head arises
on the opposite side of the humerus and is a broader and strap-
shaped muscle. It closely covers the brachialis anticus at its
1909. | ANATOMY OF CERTAIN UNGULATA, Wat
origin. In addition to these three heads there is a fourth, which
is obviously referable to the triceps complex. Its origin is from
the lower half of the humerus and its situation is posterior to and
between that of the last two heads mentioned. It is divisible
into two parallel muscles, which join before their fleshy insertion
on to the olecranon to the external side of the rest of the triceps.
Antilocapra has only three extensor muscles the tendons of
which are inserted upon the phalanges of the toes, or rather four,
of which two become united at the wrist, and may therefore be
regarded asa single muscle. In the middle of the metacarpus only
three tendons can be seen. Messrs. Windle & Parsons describe
as long extensor muscles in the Ungulata the tendons of which are
inserted upon the phalanges: (1) Extensor communis digitorum ;
(2) Extensor minimi digiti; (3) Extensor profundus digitorum,
which, however, is described as only occurring in the Elephant
and occasionally in Suidee.
The four muscles in Antilocapra have the following origins and
insertions :—
(1), (2). Two muscles arise one below (not deep of) the other,
and in close contiguity at their origin from the external condyle ;
the lower head, which is the smaller, also varies, and perhaps
may be said to chiefly arise from the shaft of the ulna quite at
the commencement of the latter. The two tendons are perfectly
distinct until a little way beyond the wrist, when they absolutely
jom. These two muscles together form, as I presume, the
extensor minime digit (or extensor digitorum lateralis).
(3). A thin muscle, arising from the external condyle under
cover of the upper part of the extensor minimi digiti, is reinforced
by a long slender muscular slip from the ulna, and ends in a
tendon which closely accompanies that of the next muscle to be
described. The tendon runs between the two toes covered by
the intrinsic extensors of the hand and supplies both digits.
This tendon with its two branches contrasts with the broad
flattened termination of the tendon of the extensor minimi
and of the next muscle to be described. The division of the
tendon into two is quite low down, in fact at the very end of the
cannon-bone. I imagine that this muscle is the extensor com-
munis digitorwn.
4). The third (see above), or perhaps fourth, long extensor
muscle of the hand of Antilocapra arises higher up the humerus,
below and in contact with the thick fleshy extensor carpi radialis.
Its tendon, which has been already referred to, supplies the
medius digit. It has no second head that I could discover.
I suppose that this muscle is the equivalent of the eatensor
profundus digitorum, which has been stated in the case of the
Pig to be inserted upon the index and medius digits *,
I now pass on to the muscles visible on the extensor side of the
forearm before considering the arrangement of the flexors.
* Hide Windle & Parsons.
Proc. Zoou. Soc.—1909, No. XII. 12
178 MR. F, E. BEDDARD ON THE [Feb. 16,
The extensores carpi radiales longior et brevior form, asin Ungu-
lates generally, a single muscle, unless indeed the muscle which
has just been referred to as the extensor profundus is really the
longior. It has the broad strap-shaped tendon described in other
forms. A muscle, which I presume to be the pronator radi teres,
Text-fig. 12.
Antilocapra americana. Musculature of outside of forearm.
a. Extensor carpi radialis. 6. Extensor profundus digitorum. . Extensor com-
munis digitorum. d & e. Two tendons just before their junction of the double
extensor minimi digiti. jf. Flexor sublimis.
1909. | ANATOMY OF CERTAIN UNGULATA. 7g)
arises from the ulna and extends over the radius for a long
distance towards the wrist.
Finally, on the extensor side of the forearm is to be noted the
extensor carpi ulnaris.
On the flexor side of the forearm the following muscles are to
be seen :-—
The first of these, beginning at the lower border, is the flexor
carpi ulnaris, which is exactly opposite to the extensor carp
ulnaris, and arises from the olecranon as well as from the internal
head of the humerus. Its insertion is parallel and contiguous to
that of the extensor carpi radialis.
The flexor sublimis digitorum arises from the internal head of
the humerus and from the surface of and in common with one of
the heads of the flexor profundus. It has only one fleshy belly,
which ends in a single tendon.
The flexor profundus digitorum is also, like the last muscle,
rather different from the corresponding muscle in other Bovide.
It has four heads. The first of them is the most superficial, and
arises from the internal condyle of the humerus in common with
the last muscle, beneath the flexor carpi ulnaris. Two heads of
unequal importance arise deep of the last and also from the
internal condyle. The fourth head is entirely ulnar in origin, in
fact from the olecranon. Its tendon joins that of the two last
described heads at about where they join each other. There is
not any muscular palmaris longus that I could find, but a very
strong tendon flattened in form and much stronger than any of
the other tendons of the hand.
The flexor carpi radialis presents no peculiarities.
I could find no lumbricales.
There were two short extensors between and below which runs
the tendon of the extensor digitorum.
Some of the muscles of the fore limb are illustrated in the
accompanying figure (text-fig. 12), and include those in which
Antilocapra certainly differs from other Ungulates according to
the elaborate account given by Messrs. Parsons & Windle of the
myology of that group.
In the hind limb, the sheet of muscle and fascia which corre-
sponds to the gluteus maximus, femoro-coccygeus, biceps, and
tensor fascie femoris is attached, as in other Ungulates, to the
fascia covering the knee and foreleg for about halfway down. Of
this mass I have not been able to recognise all the elements
mentioned above as distinct muscular sheets. The tensor fasciz
arising from the ilium in front, is the most conspicuously
separate. The others seem to me to form a continuous sheet of
muscle, arising from the tuber ischii and the caudal and sacral
vertebre in front. Three glut, besides the maximus, are easy
to dissect and separate, and are all attached to the greater
tuberosity of the femur, the gluteus medius to its free edge, and
the other two in front and near to its origin from the femur.
I have no particular remarks to offer with regard to the very
Le
180 MR. F. E. BEDDARD ON THE [ Feb. 16,
wide gracilis, the narrow sartorius, and the semitendimosus and
semimembranosus, except that the semitendinosus has only one
head from the tuber ischii. I am unable to identify with
certainty a presemimembranosus. The muscle which I term here
semimembranosus is one muscle arising from the tuber ischu and
covering over the adductor which lies deep of it. It is double
nowhere along its course, and is inserted along a semicircular
line from the end of the femur to the beginning of the tibia,
covering over the origin of the gastrocnemius. There is no
separation at the insertion into two muscles, nor is there any
rounded tendon, the insertion being mainly muscular. ‘There is,
however, some connection just at the insertion into the extremity
of the adductor and also with the head of the gastrocnemus. In
any case there is nothing like the separation of two fleshy bellies
such as are figured by Messrs. Windle & Parsons*.
With regard to the quadriceps femoris, I have only to observe
that the vastus internus is much smaller than the vastus externus.
The pyrifornis was perfectly distinct from the entogluteus.
The tibialis anticus has but one head of origin, and that is a
long flat tendon arising from the external condyle of the femur
and passing over the knee in front to form a fleshy belly which
ends in a long tendon fanning out at its insertion. A muscle
corresponding to the second (tibial) head also exists which ends
in a tendon which perforates the tendon of the last and is
attached considerably below it, 7.e¢., nearer to the ventral side of
the foot.
The extensor longus digitorum seems to me to arise only from the
fascia between the muscle itself and the femoral head of the tibialis.
Besides the tendon of this muscle another tendon runs along the
anterior face of the metatarsus. The two tendons of the extensor
longus are separate before the ankle, although they run along
the metatarsus closely bound together. The third tendon I refer
to the peroneus quarti digiti. The peroneus longus is also
present. No other peroneals were found.
I have also dissected the gastrocnemius and soleus, and find
them to be as in the Bovide. That is to say, that the small and
fleshy soleus arises from the head of the fibula and joins the
outer head of the gastrocnemius.
The plantaris is remarkable for a relationship to the gastro-
cnemius not commented upon by Messrs. Parsons and Windle.
Tn one specimen that I dissected, there is an intimate connection
between the plantaris and the outer head of the gastrocnemius
shortly after their origm. Some of the fibres of the gastro-
cnemius arise from the fascia separating it from the plantaris. In
fact, the two muscles cannot here be separated by dissection.
In the other specimen there was also a connection between the
two muscles but rather different in its kind. A slender tendon
* Loe. cit. fig. 26, p. 274, S.M. and P.S.M.
1909. | ANATOMY OF CERTAIN UNGULATA. 181
left the plantaris and swelling into a slender muscular belly
joined the gastrocnemius.
The three flexors, viz., fl. tibialis, fl. fibularis, and fl. tibialis
posticus, are present, and their tendons were as usual. It is
important to note the presence of the last-named muscle, which
has been said to be absent in Bovines.
Intestinal Tract.—Dr. Murie* says very little about the in-
testinal tract of the animal, merely giving the lengths of the several
regions and noting the absence of an ilio-cecal gland. I find
the colic spiral to be quite typically ruminant in its characters,
and to be perhaps rather large for so small an animal. In one of
the two fcetuses in which I studied it carefully, the colic spiral was
not flat, as it is insome Antelopes with but few coils, but convex.
A transverse section through the centre of the coil and including
the tract of colon immediately after it has left the cecum, would
cut through the colon fourteen times. The spiral is thus very
much longer than—for instance—that of Madoqua to be later
described. Furthermore, it is to be noted that there is in the
foetus which I examined no marked ansa paracecalis; the colon
where it emerges from the cecum is not at all bent upon itself in
the way so characteristic of many Artiodactyles and Rodents.
I could detect no definite ansa paracecalis; and in this Antilo-
capra agrees with Tragulus (see text-fig. 14, p. 184), where there
is no loop between the cecum and the spiral loop.
The cecum itself has no peculiarities; it has the usual sausage
shape and blunt tip of that of other Ruminants. There is some
variation among the Ruminants in the degree to which the ileo-
cecal mesentery extends along the cecum; in Antilocapra the
membrane extends about halfway along the cecum. The colon,
when the spiral is left, runs at first in a wide loop which is
thrown distally into a number of loose folds which are like those
of the small intestine in not being permanent. After this a short
tract of the colon is attached by membrane to the czecum; thereafter
a short and wide but fixed semicircular loop of colon ends in a
short straight rectum. I have examined and can compare with
Antilocapra as regards some of the facts just mentioned, three
species of Cephalophus, viz., C. maxwelli, C. melanorrhous, and
C. dorsalis. In all of these there is a well-marked ansa_ para-
cecalis. In none of them is there so complex a colic spiral as in
Antilocapra. Lonnberg has examined several species of Cepha-
lophust, but has not seen C’. dorsalis which presents differences
from C. maxwelli. C. dorsalis is a larger species than the latter
but is differently coloured ; it also possesses inguinal glands,
which are wanting in both of the other species I have referred
to above. While in C. melanorrhous. and C. maawelli the colic
spiral forms a complete circle and about half a circle, it forms
two complete circles and a little more in (. dorsalis. In a
* P.Z.S. 1870, p. 350. + Nova Acta Reg. Soc. Unsala, 1903.
182 MR. F, E, BEDDARD ON THE | Feb. 16,
transverse section across the coil in the two former species the
colon would be cut across eight times ; in the case of Cephalophus
dorsalis ten times. The result is of course that the apex of the
spiral is rather different in the two cases. Furthermore, while
in C. melanorrhous and C. maxwelli the ileo-ceecal mesentery is
inserted up to the very tip of the cecum, it falls considerably
short of this in C. dorsalis. .
Dr. Lonnberg* has used a very convenient method of ex-
pressing the direction of the various loops of the colic spiral,
which I adopt in comparing Anéilocapra with Cephalophus and
some other forms (see text-fig. 13). The thick black line indicates
the entering colon ; the pale line the colon which leaves the coil.
The letter X indicates a fixed point in all the figures, 7. ¢., the
extremity of the colic spiral, which is of course merely a single
loop like that of Lemur, &c., and is wrapped within its own curves
to form a spiral. It will be noticed that the simplest spiral
exhibited among these Antelopes is that of J/adoqua which will
be deseribed later. It forms a complete circle, each limb being
approximately equal. The spiral of Cephalophus dorsalis is a
further advance upon this; the spiral in this Antelope consists
of two circles, each limb again being approximately equal. It
will be noticed that in both these cases the limb of the spiral,
which is continuous with the cecal side of the colon, ends on the
same side of the spine as that on which the cecum lies, and,
furthermore, that the number of the loops of the entering and
outgoing colon are equal, 7. ¢., three of each in Madoqua and
five of each in Cephalophus dorsalis.
Tragulus stanleyanus, Cephalophus maxwelli, and Antilocapra
form another and a parallel series of ascending complexity. The
first-named is on the same level as Madoqua. The spiral has
only one turn. But the arrangement of the loops is different.
It happens that in Zragulus stanleyanus the end of the spiral
is directed away from the cecal side of the spiral, z.e., trans-
versely to the antero-posterior axis of the spiral and to the
direction of the terminal loop in the other types already described.
It will be seen, however, that if the spiral be increased by half
a turn so that the axis of the end of the spiral be made to
coincide with that of the other types, the limb of the terminal
part of the spiral which lies on the cecal side of the spiral is not
a part of the entering coil of intestine as in Madoqua and Cepha-
lophus dorsalis, but of the outgoing coil. In Cephalophus
maxwelli the spiral is more complex, 7. e. larger, but there is
precisely the same relative position of the entering and outgoing
limb of the spiral asin Zragulus. So finally in Antilocapra, which
belongs to this same series. Here, however, the tip of the spiral
is directed upwards; but if this be altered so as to make the
spiral of the same length as in Cephalophus maawelli, it will be
seen that the limb of the spiral which forms the cecal side of the
* Zoologiska Studier tillagnade Prof. T. Tullberg. Uppsala, 1907, p. 248.
1909. ] ANATOMY OF CERTAIN UNGULATA, 183
%
terminal portion is the outgoing spiral, precisely as in Cephalophus
maxwelli. It is not at all surprising-—if the facts be properly
considered—to find that Cephalophus dorsalis and C. maxwelli
are not near akin in the nature of their colic spirals. The
Text-fig. 13.
a
fa
The colic helicine in various Artiodactyles, arranged in two series, 7.e. 1, la, 18,
and 2,2a,26. The cecum is represented to the right and the ingoing limb
of the spiral is black. The outgoing limb of the spiral is left uncoloured.
1. Madoqua phillipsi. 1a. Cephalophus dorsalis. 16. Moschus moschiferus.
2. Tragulus stanleyanus. 2a. Cephalophus maxwelli. 2b. Antilocapra
americana.
differences in external characters appear to me to be quite as
great as those which distinguish certain other genera of Antelope,
184 MR. F. E. BEDDARD ON THE [Feb. 16,
and even greater than those which have been used in a few
cases. The internal difference only confirms the value of these
characters.
The spirals described in the foregoing paragraphs are com-
mitted—so to speak—to one of the two series of right-handed
spirals. There is no common basal form from which might arise
either type of spiral. Whether such exists among the Artio-
dactyla remains to be seen. It is important to notice that this
does not occur in the case of the simplest spiral known in that
nN
Text-fig. 14.
Cxeum, colic helicine, and adjacent parts of the gut of Tragulus stanleyanus.
a. Front edge of caeco-duodenal ligament. C. Caecum. col. End of colon
or beginning of rectum. AH. Colic helicine. O. Omentum.
subdivision of the Ungulata, viz., that of the Chevrotains. It does
not thus seem likely that such a spiral will be found in any genus
which has not been up to the present examined. There is no
doubt, for instance, that the Pig tribe does not help us in this
matter in the least. For in most of them at any rate, and
probably in all, the colic spiral is very complex and specialised,
and offers no hints of the primitive condition. This applies to
the Peccary as well as to the more specialised pigs. Nor do the
1909. | ANATOMY OF CERTAIN UNGULATA. 185
remaining subdivisions of the Ungulates help us in this search.
In neither Hyraxz nor Hlephas are there any beginnings of the
spiral. It is true that the simple colic loop of the Perissodactyles
(found in all the families of Perissodactyla) may be regarded as
the progenitor of the spiral; but this is going too far back in the
evolution of the spiral. What is wanted is a simple spiral of a
negative character which with further growth might evolve into
either type. Now it is interesting to find that this form of spiral
—precisely what is wanted to serve as a starting point—is to be
seen in those Lemurs with a simple colic spiral, i. e., Galago*,
Loris, Perodicticus, and Nycticebus. 1t will be seen from an in-
spection of the figure that this simple spiral can be easily regarded
as a three quarters of a circle spiral with the apex of the spiral
pointing upwards, and of course with the outer limb of the
descending part of the colon turned towards the cecum. On the
other hand, it may be regarded as a commencing spiral of half
a turn with the apex pointing downwards. In this case it
obviously belongs to the other series, for if the spiral were
continued so that the apex pointed upwards, the limb forming
the apex and nearest to the cecum would be that of the inner
limb of the loop. It is perfectly clear, therefore, that a growth
of this rudimentary spiral could produce either of the two types
of Artiodactyle spiral with which I am concerned in the present
communication. This is of course not tantamount to insisting
upon any close relationship between the Ungulates or the Artio-
dactyle Ungulates and the Lemuroideay. But it is at least
important to note that while the only widely distributed sub-
families of the Lemuroidea have preserved without variation an
archaic and indifferent form of spiral, the most archaic Artio-
dactyles have not the simplest form of spiral conceivable. So
far, therefore, it would appear as if the colic spiral as such had
been inherited by the Artiodactyles. It is, of course, not at all
certain that the single loop of the Perissodactyle Ungulates is
the homologue of the Artiodactyle spiral; it may represent the
ansa paraceecalis of the latter.
The facts detailed under the above descriptions of the anatomy
of Antilocapra may be supplemented by some account of the
intestinal tract in other Artiodactyles which I have had recently
the opportunity of examining at the Prosectorium. Among
those the most interesting perhaps is that of the Musk Deer
(Moschus moschiferus) . The viscera of this animal, including
the alimentary tract, have been described by Sir W. H. Flower §
and Prof. Garrod ||. The cecum is figured by Flower, but the
colon is not specially described. A footnote adds that in
* For non-diagrammatic figure see Beddard, P. Z. S. 1908, p. 578, fig. 114.
+ Moreover, the apparently similar colic spiral found in certain Rodents (Hydio-
cherus, Dasyprocta, and Cologenys) requires further consideration trom this point
of view.
+t A female dissected Dec. 21, 1908.
§ P. Z. S. 1875, p. 159. || P. Z. S. 1877, p. 287.
186 MR. F. E. BEDDARD ON THE [ Feb. 16,
the Pudu Deer (Pudua humilis), of which a specimen was
dissected at the same time by Sir W. H. Flower for the purpose
of comparing with JMoschus, a deer of about the same size, ‘“ the
colon... . forms a very simple helicine coil, composed of two
complete circles in one direction and of two in the other” *.
It is not clear whether it is to be inferred that JJoschus agrees.
In any case, Garrod describes the colic spiral as having three and
a half double turns. This statement is approximately correct, as
Text-fig. 15.
et EI
Czecum, colic helicine, and adjacent parts of the gut of Moschus moschiferus.
p. Ansa paracecalis. Other lettering as in text-fig. 14.
will be shown by the annexed figure (text-fig. 15), which gives
the same number of coils as has the spiral of Antilocapra. The
termination of the spiral, however, points downwards, in which
Moschus disagrees with Antilocapra. Nevertheless, it belongs to
the same series as that which commences with J/adoqua (see
183). With regard to the ileo-cecal ligament I confirm
Sir W. H. Flower that it is not attached up to the very end of the
23 ILC, Cote WW LAL
1909. | ANATOMY OF CERTAIN UNGULATA. 187
cecum. It is important to note that this character is apparently
a constant one, which I have used above (p. 181) in distinguishing
other Antelopes. The ansa paracecalis of Moschus is not dealt
with by either Flower or Garrod. It is remarkable in that it is
bent upon itself four times to form a small spiral, which is quite
as conspicuous a spival as is that of the Lemurine subfamilies
Galagine and Lorisine, and may even, quite possibly, really be
the equivalent of that colic loop—the true ansa coli dextra of the
Rodents being in that case absent from the Lemurs.
In Gazella rufifrons (a female) the ansa paracecalis is roughly
of the shape of the letter “‘m.” That is to say, it consists of two
parallel loops. Immediately after the ansa paracecalis the colon
diminishes suddenly in calibre, and enters the region of the
colic spiral or ansa coli dextra. This is a comparatively simple
coil. It is only a little more complicated than that of JMZadoqua
described on page 189. Cut right across, the colon would be
divided seven times, counting the straight tracts which enter and
which leave the coil. As usual, the small intestine is festooned
along the colon after the colic spiral. The colon in this region
passes almost completely round the circumference of the abdominal
cavity, and forms nowhere any special loops. The duodenum where
it turns upon itself is fixed, as in other Antelopes, to the colon
itself, and not to the mesocolon
With regard to the colic spiral, it is to be noted that it belongs
to the type of Cephalophus ma: gwelli. Bub the loops do not form
a regular watchspring as in those Artiodactyles already con-
sidered. For the terminal loop of the spiral does not lie entirely
within the coil but extends beyond it.
Dr. Lonnberg comments upon the remarkable colic spiral of
the Musk-Ox*. Im addition to the spiral coil there is a
separate loop formed by the outgoing limb of the spiral, as is
depicted in his figure. It seems to me that this state of affairs
is comparable to the presence in so many Rodents of two anse
coli, the dextra and sinistra. The two are close together in
Squirrels, for example. If one of these—and they are frequently
of different sizes—were converted into a spiral, we should have
a condition produced which would be precisely like the Musk-Ox.
For the lengthening of the one loop—already the longer of the
two-—to form a spiral would bring it about that the shorter ansa,
remaining a simple tubular loop, would be affixed to the back or
to the front of the spiral, as is actually the case in the Musk-Ox.
There is, however, no need to theorise ; for the Rodent Dasyprocta
punctata shows exactly the same conditions as those met with in
the Musk-Ox. In this species * there is a short spiral and also a
simple loop closely affixed to it. I take it that this is a simple
form of the complex spiral and associated Hore} found in the
Musk- Ox:
But woaly in one specimen out of two; the other showed a single spiral only.
PZ
aK
+ P. Z. 8, 1900, p. 152, fig. 7.
188 MR. F. E. BEDDARD ON THE [ Feb. 16,
5) Some Notes unon the Anatomy of Madoqua phillipsi.
ip y quay p
A dissection of an example of this small Antelope enables me
to make a contribution to the knowledge of the structure of the
Cavicorn Ungulates, for I believe that we have at present no
information as to the anatomy of this pentigmbn genus of
Antelopes, beyond the notes contained in Lénnberg’s paper already
referred to.
The example which I examined was a female. The usual
four teats were present and also as usual the anterior pair were
much further away from each other than the pair lying behind
them. There was no trace that I could see of the inguinal
glands, and, indeed, the inguinal region, instead of being bare,
was particularly well covered with long hair. I have notes
upon some of the organs, and I naturally paid special attention
to those structures which are known to vary in these animals.
As to the Vascular System I found that the heart shows the
“‘moderator band” so frequently found in running animals—
birds as well as mammals*. It was a considerable fleshy
column connecting together the free and septal walls of the
right ventricle. The postcaval vein was typical in its arrange-
ment, 7.e. the vein lay postrenally to the right of the aorta.
The azygos had the following structure :—
The general plan is like that of other Artiodactyles. That is
to say, “the ri ight azygos is much less important than the left.
The latter opens moreover, as is usual, directly into the heart,
but it consists of two portions, an anterior and a posterior.
They unite just opposite to the transversely running vessel
which is formed by their union and debouches into the heart.
There is thus, as I may point out, a remnant of the left anterior
cardinal, which I presume the anterior branch of the left azygos
to represent. This fact in the structure of the azygos of
Madoqua is furthermore of some systematic interest. For, as 1
have discovered tT in Rhaphicerus and Ourebia, the arrangement
of the veins bringing back blood from the dorsal parietes on the
left is quite like that which has just been described in J/adoqua.
Now these three genera have been placed in the same subfamily
Neotragine. This additional structural likeness justifies still
further this placing. The right azygos of J/adoqua is but feebly
developed and brings back blood from only two or three inter-
costal spaces.
In the intestinal tract of Antelopes it is the colon which
shows the principal differences. I have therefore made careful
notes upon certain points in the structure of that part of the gut
in Madoqua phillipsi. The cecum is long and like that of
other Antelopes in its blunt ending without any diminution
of calibre. The iewm enters it at an acute angle and is attached
to it hy mesentery which extends a long way towards the tip of
* To which particular attention was called by the late Prof. Rolleston.
+ “On the Azygos Vein in the Mammalia,” P. Z.S. 1907, pp. 184, 185.
1909. ] ANATOMY OF CERTAIN UNGULATA. 189
the cecum. The colon as it leaves the cecum shows hardly any
trace of the paracecal flexure so common in these animals.
There is practically no twisted section of colon at this point.
Furthermore, the colic spiral is very much reduced, a fact which
one naturally associates with the small size of theanimal. It isin
fact, as the accompanying text-figure shows (text-fig. 13, p. 183),
no more complicated than in the Lorisine Lemurs*, which show
the simplest colic spiral known to me. A transverse section
across this spiral would in fact only divide the colon five times.
This of course recalls the diminutive spiral of the Chevrotains
described by Milne-Edwardst and more recently by Mitchell $.
It is to be noted, however, that the spiral in Zragulus (see text-
fig. 14, p. 184) is rather different from that of Jadogua. For in
the latter the spiral is flat, and is separated from the cecum by
the ileum to which it is attached by mesentery, just as the ileum
in its turn is attached to the cecum by a band of mesentery. In
Tragulus the small spiral is fixed on to the cecum near to its
colic end. It appears to me possible that this spiral in 7ragulus
is not the precise equivalent of that of other Artiodactyles, but is
the ansa paracecalis which tends to show a spiral form in many.
For example, in the small Antelope Cephalophus abyssinicus, of
which I have lately dissected an example, the ansa paraceecalis was
much more conspicuous and complicated than that of Madoqua
and tended towards a spiral arrangement; so too with Moschus
described and figured above. It may also be that the distinctly
spiral commencement of the ansa paracecalis in Hyrax§ is
referable to the same category, as is also that of many Rodents.
There is, however, some variation in the colic spiral of
Ruminants which may be a question of age or of actual variation.
I have pointed out that in the Rodent Hydrochawrus|| the younger
individuals have a rudimentary spiral which later becomes large.
I have examined this spiral in two examples of the Antelope
Tragelaphus scriptus, both females. In one specimen the coils
were hardly more complex than in Madoqua. The colon consisted
in this region of four loops each consisting of two limbs, so that a
transverse section across this region of the gut would divide the
colon eight times. On the other hand, the second specimen had
a more complex colic spiral.
I may observe that the close connection of the small intestine
with the colon—for the former gut is attached by mesentery to
the colon for some distance after the colon has emerged from the
colic spiral—is a feature often (? always) found among Artio-
dactyles; it is paralleled in a remarkable way in the Beaver,
another fact illustrative of the numerous structural resemblances
between the Rodents and Ungulates.
* Beddard, P. Z.S. 1908, p. 578, and literature therein cited.
+ Ann. Sci. Nat. 1864.
t Trans. Zool. Soc. vol. xvii. p. 472, fig. 19.
§ P.Z.S. 1908, p. 532.
|| P.Z.S. 1908, p. 537.
190 MR. F. E. BEDDARD ON THE [ Feb. 16,
Dr. Linnberg has specially devoted himself to the intestinal
tract (and other features in the anatomy) of the Artiodactyles ”.
As these animals are comparatively little known in the anatomy
of their soft parts, Ido not hesitate to set down some further
facts as a contribution to our gradually increasing knowledge of
the group, which may prove to be distinctive not of the particular
species now under consideration but of the group generally. In
Madoqua, as in Cephalophus abyssinicus, the end of the duodenal
loop is not fixed by the ligamentum cavo-duodenale to the post-
caval vein or to the mesocolon, but to the ventral surface of the
colon itself.
Some of the recto-duodenal ligamentum strays over on to the
adjacent right kidney. I could find no definite hepato-renal
ligament. But the right extremity of the liver was attached to
the parietes by a well-marked ligament. I may remark also that
in Cephalophus abyssinicus the duodenum was attached to the
rectum itself in precisely the same way by a ligament, and that
there was also the same attachment to the right kidney. The
left kidney was situated a long way below the right kidney,
there being a considerable distance between the posterior end of
the right kidney and the anterior end of the left kidney. The
spleen of Madoquwa had the more or less circular contour of this
‘“oland” in some other Ruminants. It was as firmly fixed to the
stomach throughout as in other Ruminants. There was in fact
no free gastrosplenic omentum.
The Brain of Madoqua phillipsi is not known, so far as I am
aware, and will serve to illustrate the brain characteristics of a
small Antelope with the suici in a simple condition. It may be
compared with that of Antiocapra, which I also figure in the
present communication. The general outline seems to me to be
remarkable for the square outline and termination anteriorly of
the portion of the brain lying in front of the Sylvian fissure.
This form of brain is still more marked in the Pronghorn, which
T have dealt with on a previous page. Secondly, as will be seen
in the drawing of the dorsal surface submitted herewith (text-
fig. 16), the corpora quadrigemina are partially visible, which is at
least unusual among the Ungulata. The third obvious peculiarity,
also to be seen in the same dorsal view, is the exposure of the
splenial suleus which runs on each side a curved course at some
little distance from the median fissure of the brain. The exposure
of this fissure is not unknown in other Artiodactyles—it occurs,
for instance, in Vragulust, Dorcatheriwm, Moschus, Cervulus—
but it must not be confused with an entolateral sulcus, such as
occurs in Cervus damat, and which at first sight might be
confused with it. Its continuation with the crucial anteriorly
* Nova Acta Reg. Soc. Upsala, 1903; K. Vet.-Ak. Handl. 1901; P.Z.S. 1900,
p. 142; and in Zoologiska Studier tillagnade Professor T. Tullberg, 1907, p. 237.
+ See Cat. Physiol. Series, Roy. Coll. Surgeons, Vol. 11. 2nd ed., 1902, p. 323,
p. 324, p. 325 fig. 187, p. 327.
+ Ibid. p. 329, fig. 191.
1909.] ANATOMY OF CERTAIN UNGULATA. 191
shows that in Wadoqua the fissure in question is really a splenial.
Dr. Elliot Smith has remarked * that “the position of this sulcus
in the Ungulata seems to be determined largely by the size of
the hemispheres,” it being visible above in Zragulus and not in
Cervus. So far as Madoqua is concerned this generalisation
would seem to be true of the Cavicornia as well as of the other
Ruminants. The brain of the Sheep has been so often described
and figured 7 that it may be well regarded as a convenient basis
of comparison for other hollow-horned Ruminants. There are in
Madoqua, as in Ovis, three lateral fissures of which I identify
the middle one and the longest as the lateral, the other two
being ecto- and ento-lateral respectively. The lateral bends
outwards anteriorly, but does not end in any other Sssure.
Text-fig. 16.
Brain of Madoqua phillipsi, dorsal view. Natural size.
a. Coronal suleus. _c. Inner branch of the same. d. Suprasylvian sulcus.
e. Lateral sulcus. f Entolateral sulcus. g. Splenial sulcus.
The suprasylvian fissure is the same on the two sides of the
body and consists of precisely the same elements in the Sheep,
excepting that I did not find the fissure lettered by Holl £ “rdss”
and represented in the drawing given by Elliot Smith not far
behind the Sylvian fissure, but not lettered. The suprasylvian
fissure, however, bifurcates posteriorly in a V-shaped fashion as
indicated by Holl in Ovis; but in Madoqua the two limbs are
longer, the downwards directed limb running parallel with the
suleus obliquus. The anteriorly situated “inferior” branch of
the suprasylvian joins the sulcus coronalis.
%* Tbid. p. 326. ost
+ Most recently by Elliot Smith (loc. cit. p. 340, figs. 194, 195, 196) and Holl,
Arch. f. Anat. 1900, pl. xvii. figs. 7, 8, 9.
{ Loc. cit. pl. xvi. fig. 7.
192 MR. F. E. BEDDARD ON THE [ Feb. 16,
The Sylvian fissure in the usual Artiodactyle fashion starts
from the edge of the depressed gyrus arcuatus. On one side of
the body it is short and runs in a straight course dorsally, being
just hooked over anteriorly at the summit. On the other side it
is shorter still. There is nothing particular to say of the anterior
and posterior ectosylvians.
Text-fig. 17.
Brain of Madoqua phillipsi, lateral view. Natural size.
s. Sylvian fissure. Other letters as in text-fig. 16.
The coronal sulcus, whose course on either side is straight
and parallel with the median sulcus, sends a branch inwards
at its posterior end as in other Artiodactyles. This branch,
however, does not reach the middle line of the brain on either
side. In front of this the gyrus lymg between the median sulcus
of the hemispheres and the coronal sulcus is indented by the
emergence of the splenial sulcus, this indentation being, as I
presume, the equivalent of the crucial sulcus in other <Artio-
dactyles such as Ovis*.
(6) The Brain of Babyrussa alfurus.
Although the brain of this Suid has been already described by
Vrolik +, the description and figures were published a great many
years ago and, moreover, do not agree in every detail with the
specimens which I have myself had the opportunity of studying.
T have examined carefully the brains of two examples of Labyrussa
alfurus which have died in the Society’s Gardens during the last
four yearst, one of which has been drawn for me by Mr. W.S.
Berridge, F.Z.S. and is represented in the accompanying text-
figure (text-fig. 18, p. 194). Theanimal wasa female. The length
of the hemispheres in a straight line is 82 mm., the greatest
diameter of the brain 62 mm. I have compared this brain with
* Blliot Smith, Joc. cit. fig. 195, p. 340.
+ Vrolik in N. Verhand. Nederl. Inst. Amsterdam, vol. x. 1844, p. 207. The
paper is written in French. The brain is figured in dorsal and ventral views and
in section.
t The first died June 29th, 1905, the second Jan. 6th, 1908.
1909. | ANATOMY OF CERTAIN UNGULATA. 193
specimens of Sus cristates and Dicotyles tajagu, of both of which
brains exist in the collection at the Prosectorium.
The coronal sulci are much more obliquely set than in either
Sus or Dicotyles, especially more so than in Dicotyles, where the
furrows in question, save just where they dip into the middle
line, are straight and parallel with the long axis of the brain.
The brain of Babyrussa in fact agrees in this particular with
that of Phacocherus ethiopicus figured by Elliot Smith *. Not
so, however, in their proportionate length; for while in Phaco-
cherus—judging from the figure already referred to—the coronal
suture of each side is very nearly half the length of the brain,
the length of each of these sulci in Babyrussa is 37 mm., measured
from the anterior end of the brain to the point where they meet
in the middle line of the brain; the rest of the brain measures
about 55mm. ‘These measurements are taken along the curvature
of the brain, and are therefore in excess of that just given of the
length of the hemispheres.
In Dicotyles tajacu the proportions are reversed, the anterior
part of the brain to the posterior end of the coronary sulci is
actually longer than the rest of the hemispheres. In Svs,
however, the proportions would appear to be very much as in
Babyrussa, and probably by making allowances for the curvature
not measurable in the figure quoted, Phacochwrus is not very
different. Elliot Smith agrees with Garrod? in regarding as
distinctive of the Pig tribe the blending of the coronal and
interealary (or splenial) sulci as compared with other Artio-
dactyles.
The intercalary or splenial sulcus in Babyrussa (which obviously
suggests—if it be not comparable to—the calloso-marginal of
Apes) is continuous with the genwal sulcus in front, as in Sus
and Dicotyles. It is also continuous with the coronal, bifurcating,
in fact, as it were anteriorly to form the coronal and genual. In
this Babyrussa of course agrees with other Pigs, as has been
already mentioned. It does not, however, agree with an example
of Dicotyles tajacu which I have in my possession. I am not
quite able to understand precisely what the late Prof. Garrod
meant when he wrote concerning the splenial fissure of Dicoty/es
as follows £:—“ There is one upward branch of the splenial fissure
which joins § the fissura coronalis, and is not a continuation of it,
as in Sus.” Dr. Elliot Smith in dealing with the brain of the
same species repeats Garrod’s word “joined.” I find in the only
brain which I have exainined no communication whatever between
“the cingular are of fused calcarine, intercalary and genual sulci”
and the coronal sulcus on either side of the body. <A bridging
convolution occupies the place where this fissure would otherwise
lie. But if the coronal sulcus were continued back to join the
* Cat. Physiol. Series, Roy. Coll. Surgeons, vol. 11. 2nd ed. 1902, p. 316, fig. 182.
+ “On the Brain of Hippopotamus,” Trans. Zool. Soc. x1. 1880, p. 13.
i Hoc. cit. p. 14.
§ The italics are Mr. Garrod’s.
PRoceZo0oLy SOU.) 909s Non xelule 13
194 MR. F. E. BEDDARD ON THE [ Feb. 16,
splenial (and the gap separating them is but small) there would
be no difference in the mode of their connection from that
shown in the brain of the Babyrussa. There would be a similar
appearance of the bifurcation of the splenial that has been
yeterred to.
Text-fig, 18.
Brain of Babyrussa alfurus, dorsal view. ‘Three-quarters natural size.
a. Coronal suleus. 6, ec. Right and left branches of splenial. d. Suprasylvian
sulcus. e. Lateral sulcus. f. Entolateral sulcus.
Neither Mr. Garrod nor Dr, Elliot Smith mentions the interesting
fact that the convolution lying between the splenial fissure and
the corpus callosum (7. e. the hippocampal gyrus ?) appears on the
surface of the brain and is—posteriorly at any rate—not much
depressed below the level of adjoining regions of the brain in the
Peceary, while it is completely concealed in the Babyrussa as in
Sus, and apparently in Phacocherus. 'This arrangement is seen
in other Ungulates besides Dicotyles. In Babyrussa the splenial
gives rise to another fissure lying behind the oblique furrow
which joins it with the superficially running coronal. This is
1909. ] ANATOMY OF CERTAIN UNGULATA. 195
also well known to occur in Sus. But in the specimen of
S. eristatus which I examined the fissure did not reach the
splenial on either side. It may be that this fissure is the real
equivalent of what I have described above in Dicotyles as the
corono-splenial junction. The fissure mentioned by Garrod as
the equivalent of the latter does not, however, exist in my
specimen.
In any case it is clear that Babyrussa agrees with the Old World
Pigs, and differs from Dicot yles, in the shorter and more obliquely
running coronal fissures.
The lateral fissure of Babyrussa is less oblique in direction than
that of Sus, where the two furrows converge greatly anteriorly.
There is a well-marked entolateral fissure, and indeed on the
right side of one of my two specimens a second entolateral within
the first. There is, furthermore, in Babyrussa as in Sus an
ectolateral fissure, which is wanting—as is also the entolateral—
in Phacocherus* and in Dicotyles. In both specimens of the brain
of the Babyrussa which I possess the ectolateral fissure tends to
be rather imperfect, being interrupted along its course by bridging
convolutions, and is better developed anteriorly than posterior ly.
Text-fig. 19.
Brain of Babyrussa alfurus, lateral view. One-half natural size.
s. Sylvian fissure. d. Suprasylvian.
In Sus, on the other hand tf, the posterior part of this sulcus is
the most clearly marked, as was the case on one side (the right)
of one of the Babyrussa brains. It is to be noted that this
ectolateral sulcus gives off many transverse branchlets in Baby-
russé as in Sus; it is these small transverse furrows only which
are visible in Dicotyles.
The suprasylvian fissure is much like that of Sus; and I could
see no variations of moment in the two brains of Labyrussa nor
on the two sides of each brain. The fissure curves round
posteriorly towards the ventral surface of the brain, this part
corresponding, according to Garrod, with the descending limb of
the same fissure in other Artiodactyles. I may remark that
Babyrussa also agrees with the Wild Swine (Sus cristatus) in
possessing a small descending limb of the infrasylvian in front of
* Elliot Smith, Cat. Roy. Coll. Surgeons, f. ¢. fig. 182, p. 316.
Owen, ‘The Anatomy of Vertebrates,’ vol. iii. 1868, p. 123, fig. 104.
196 MR. F. E, BEDDARD ON THE [ Feb. 16,
the ascending limb of the same, which, as already described, is
connected with the splenial. I do not find this fissure in
Dicotyles.
It does not seem to me that there can be any doubt as to what
is to be regarded as the Sylvian fissure in Babyrussa. Both
specimens agree entirely in the position of the fissure which I
thus identify ; the only differences were slight ones of length and
direction of curvatures. I consider as Sylvian the fissure which
arises, as Shown in the accompanying figure (text-fig. 19, p. 195),
from the point where the nonpallial portion of the hemisphere is
widest, where the rhinal fissure, that is to say, is most convex
towards the pallimm. But I am not to be understood to deny
that this may be a ‘ pseudo-sylvian’ fissure formed by the meeting
of two lips over # true Sylvian fissure. Deep down between the
opercula an island of Reil is visible, precisely as Dr. Garson has
figured* in the Pygmy Hog (Sus salvania). The figures of the
brain of the Babyrussa which are appended to this paper show,
in fact, that the characters of the brain are those of other Suide,
and that the divergences from the prevailing pattern in that
group are only slight, and not at all in the direction of the New
World Dicotyles, nor of the remaining allied family, that of the
Hippopotamuide.
Résumé of New Facts.
The following are the principal new facts contained in the
foregoing pages and stated as briefly as possible, with references
to the pages where the full description will be found.
(1) The Himalayan Tapir (Vapirus indicus or malayanus), like
the two species of Elephant, has a pleural cavity which is
mainly obliterated by the formation of a dense network of
strands of connective tissue uniting the two layers of the
pleura.) (Seep. L613)
(2) Hyrax dorsalis like H. syriaca (described by Lonsky) has a
third ceecam about an inch long lying between the unpaired
and the paired ceca. The orifice of the ileum into the
unpaired cecum has, unlike what is found in H. syriaca,
a circular valve. There are specific differences between
HI. dorsalis and H. capensis in the cecum. (See p. 166.)
(3) The azygos vein in Hyrax capensis is, as a rule, confined to
the right side of the body. An additional vein on the left
side is rare:. (See p. 162.)
(4) The sulci of the brain of Hyrax capensis + show considerable
variation. This is especially noticeable in the Sylvian and
the pre- and post-Sylvian. (See p. 163.)
(5) Hyrax agrees with Hlephas in possessing a free fold of
peritoneum of considerable dimensions attached to each testis
* Toc. cit. (on p. 170).
+ This is not an entirely new fact, I extend the data of Dr. Elliot Smith.
S108) ANATOMY OF CERTAIN UNGULATA. 197
and floating in the abdominal cavity, the homology of which
is doubtful. (See p. 168.)
(6) The posterior region of the csophagus in Sus (Porcula)
salvania is covered by a sheet of muscle which arises as a
stout muscular slip from the vertebral column from the
centrum of the 8th or a closely neighbouring vertebra.
This muscle has not been met with in other mammals, but
may correspond to minute fascicles which in man bind down
the esophagus to adjacent membranes massed into one large
muscle. (See p. 170.)
(7) The musculature of Antilocapra americana shows certain
differences from that of other Bovines in the extensors and
flexors of the arm and leg. (See p. 175.)
(8) The colic helicines of Antilocapra, Cephalophus, Madoqua,
Moschus, Gazella, and Tragulus are shown to be referable to
two categories according to whether the emergent spiral
limb of the colon lies to the right or to the left of the
entering spiral. Even the two simplest spirals known among
Artiodactyles, i.e. those of Madoqua and Tragulus, conform
to the one or to the other plan. (See p. 182.)
(9) Although two species alleged to be of the same genus, viz.
Cephalophus dorsalis and C. maawwelli, show different spirals,
it is by no means certain that these two species are rightly
referred to one and the same genus. (See p. 181.)
(10) In no Artiodactyle (whose anatomy is known) is the common
form, from which the two types may have diverged, of spiral
visible. But the rudimentary spiral of the Galagine and
Lorisinze (among Lemurs) is a spiral from which both types
of Artiodactyle spiral might be derived. (See p. 185.)
(11) The remarkable spiral of the Musk-Ox, described by
Lénnberg, in which the emergent limb is bent once upon
itself before leaving the region of the spiral, is closely
paralleled in one specimen (out of two) of the Rodent
Dasyprocta punctata, (See p. 187.)
(12) The azygos vein of Madoqua is like that of Rhaphicerus and
Ourebia in that the left azygos has an anteriorly running
branch as well as the chief posteriorly running trunk. These
genera are usually placed in the same subfamily—Neotragine.
(See p. 188.)
(13) In the brain of Madogqua, as in that of other small
Ruminants, the splenial sulcus is exposed and superficial in
position. (See p. 190.)
(14) The brain of Babyrussa is like that of other Old World
Suide. Its slight divergences are not in the direction of
the New World Dicotyles. (See p. 192.)
(15) The brain of Antilocapra is remarkable for its squared-off
ending anteriorly and for the fact that, as in the Suide, the
crucial and coronal sulci are confluent. (See p. 173.)
198 DR. E. L. TROUESSART ON THE [Feb. 16,
3. Le Rhinocéros blane du Soudan (Rhinoceros simus cotton: ).
Par le Dr. E. L. Trovessart, C.M.Z.S., Professeur au
Muséum d’ Histoire Naturelle de Paris.
{ Received January 21, 1999. |
'PlatesXXIX.-XXXI.]
Bien que les cornes du Rhinoceros simus soient, depuis le
Moyen-Age, Vobjet d'un commerce important dans le Soudan,
et bien que plusieurs Musées d’Europe possédent depuis long-
temps de ces cornes, eest seulement en 1900 qu'un crane,
rapporté par le Major A. Gibbons de Venclave de Lado, permit
Waflirmer avec certitude que cette espéce, en voie (extinction
au sud du Zambéze, était représentée, dans la région qui s‘étend
entre le Haut-Nil et le lac Tchad, par une sous-espéce distincte.
Depuis cette ¢époque, d’autres spécimens, rapportés par le
Major P. H. G. Powell-Cotton de la méme localité (Lado),
ont permis & M. R. Lydekker de caractériser cette sous-espece
sous le nom de Rhinoceros simus cottoni*.
Il ne semble pas que cette remarquable sous-espéce ait encore
été figurée d’une maniére satisfaisante. Je saisis done Poccasion
de donner ici, de profil et de face, la photographie d'un beau male,
tué en 1908, dans le Bahr-el-Gazal, par un sportsman qui desire
garder Vanonyme. La Planche X XIX. a été exécutée d’aprés ces
photographies, et la figure de la femelle Vaprés une corne appar-
tenant aux collections du Muséum d'Histoire Naturelle de Paris.
Si la figure du Rhinoceros simus de Afrique australe, publiée
en 1894 par M. Coryndont dans les ‘ Proceedings,’ d'apres le
spécimen monté du Musée de Tring, était exacte, le Rhinoceros
simus cottoni différerait trés notablement du type du Mashona-
land. Ce dernier, 4 part le caractére du museau, est figure
comme ayant les mémes proportions que le Rhinoceros bicornis.
Le Rhinocéros blane du Soudan a des proportions bien différentes
et que l’on pourrait qualifier de “plus élégantes,” si cette
expression pouvait s’appliquer a un Rhinocéros. Le corps est
plus élevé et plus court, le garrot forme une saillie trés notable,
et par contre la croupe est déclive, wvalée suivant le terme
consacré en hippologie. Chez le Rh. bicornis, le garrot n’est pas
plus élevé que la croupe.
Mais un caractére qui est nettement accusé sur nos deux
photographies, c’est que la peau porte des tubercules arrondis
et saillants, qui rapellent ceux des Rhinocéros asiatiques, bien
que moins développés. Ce caractére parait faire défaut au
Rhinoceros simus de V Afrique australe aussi bien qu’au Rh. bi-
cornis, dont la peau ne présente que des plis assez irréguliers.
Lceil parait aussi plus développé ici, si Jen juge par un dessin de
* R. Lydekker, ‘Field, 22 February, 1908, p. 319; id. ‘The Game Animals of
Africa,’ 1908, p. 38.
+ Proce. Zool. Soc. 1894, p. 329, pl. 18.—La figure donnée par A. Smith, Ilustr.
of Zool. South Africa, 1849, pl. 19, semble plus exacte.
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la téte du RA. sinvws que j’ai sous Jes yeux et qui porte la légende :
“ Dessiné dans TAfrique Méridionale daprées la nature par
Wan J. Burchell” *.
D’aprés les renseignements que l’on posséde actuellement sur la
distribution géographique de la forme septentrionale de Vespéce,
elle existe non seulement dans l’enclave de Lado, mais aussi dans
le Bahr-el-Gazal et le Wadai, probablement jusqu’au lae Tchad.
La question de savoir si lespece existe plus au sud, entre le
Zambeze et les sources du Nil, notamment dans la région des
Grands Lacs, doit etre réservée. En effet, Speke, au cours de
son voyage de Zanzibar a Uganda (en 1860), parle de Rhinoceros
“blanes et noirs” rencontrés dans le Karagwe, par conséquent
entre les lacs Tanganyika et Kivu et le Victoria Nyanzat. Ce
serait méme la, si Je ne me trompe, la plus ancienne mention,
faite par un voyageur digne de foi, de la présence du Rhinoceros
simus au nord du Zambeze. Les trés longues cornes décrites
sous les noms de 2h. oswelli Gray = et Rh. holmwoodi Sclater §
doivent étre rapportées au male ou a la femelle de la présente
espece, et la provenance de celles-ci (par Zanzibar) indiquerait que
Vanimal qui les porte se trouve sur la rive gauche du Zambeze.
Mais, on peut aftirmer que la présence de cet animal dans
le nord de l'Afrique était déja connue dans l’antiquité grecque et
romaine. Diodore de Sicile, contemporain de Jules César, parle
dun Rhinocéros d’ Ethiopie qui portait “a Vextrémité des narines
une seule corne un peu aplatie et presquaussi dure que du fer,”
et qui livrait combat al’Eléphant. Cette corne “ un peuaplatie,”
placée a l’extrémite des narines et, de plus, suffisamment aigué
pour qu’on puisse supposer, a tort ou a raison, qu'elle est capable de
percer le ventre de / Eléphant, ne peut s’appliquer qu’au RA. simus.
Cette espece ayant en réalité deux cornes, on a longtemps mis
en doute l’exactitude de ce passage de Diodore. Mais Fresnel),
en 1848, nous apprend que les Arabes, qui font commerce des
cornes du Fh. simus, sont absolument persuadés que animal n’a
quune seule corne. La corne postérieure, toujours beaucoup
plus petite, souvent tout-a-fait rudimentaire chez la femelle, est
probablement négligée par le commerce, ou confondue avec les
cornes du 2h. bicornis, considérées comme de qualité inférieure.
Les Arabes du Soudan distinguent a premiere vue les cornes des
deux espéces. Le commercant Arabe de Djeddah qui renseignait
Fresnel sur ce prétendu Rhinocéros unicorne, lui en donna la
preuve. Il fit sortir dans la rue un serviteur qui portait une de
ces grandes cornes sur chaque bras, et Fresnel entendit les passants
prononcer, sans hésiter, le nom d’Abou-Karn (possesseur d’une
corne), tandis que le Rhinoceros bicorne ordinaire s’appelle Ahertit.
* Ce dessin est reproduit dans le‘ Bulletin de Ja Société Philomatique,’ 1817, pl. en
face de la p. 100.
+ Speke, ‘ Journal of the Discovery of the Source of the Nile,’ 1863, p. 197.
t Proc. Zool. Soc. 1854, p. 46, fig. 1.
§ Proc. Zool. Soc. 1893, p. 517, figs. 1 et 2.
|| Fresnel, “Sur Vexistence dune espéce unicorne de Rhinocéros dans la partie
tropicale de l’Atrique,” Comptes-Rendus de PAcad. des Sciences de Paris, 1848,
in SOM a Asill,
200 ON THE WHITE RHINOCEROS OF THE SOUDAN. [ Feb. 16,
Les négres du Soudan s’emparent du Rhinoceros simus en
creusant de grandes fosses, masquées par de la terre et du
feuillage, sur le chemin qu'il a coutume de suivre. Contraire-
ment a l’opinion des voyageurs dans le sud de l’Afrique, on
considére, dans le Soudan, la grande espece comme plus dan-
gereuse que le Rh. bicornis ordinaire. Le voyageur Tunisien
Mohammed-al-Tounisi, qui visita le Wadai vers 1825 *, rapporte
que le Sultan de ce pays fut chargé par un Rhinocéros blanc.
Nous savons aussi que, dans le sud de l'Afrique, Oswell eut son
cheval éventré sous lui par un 7h. simus.
Le commerce considérable de cornes de Rhinocéros des deux
espéces d’Afrique que les Arabes font, depuis une époque reculée,
semble peu connu en Europe. Ces cornes sont exportées, par les
ports de la Mer Rouge et de l’Océan Indien, pour lAvabie,
la Perse et la Chine, ot on les sculpte comme de livoire. On en
fait des coupes, des manches de couteaux, de poignards et de
sabres, des plaques de ceintures et d’autres objets. Le Muséum
d’ Histoire Naturelle de Paris possede trois coupes sculptées dans
de la corne de Rhinocéros; lune delle est dun travail tres
artistique et d’un gout trés pur, que ne renierait pas un sculpteur
européen. La corne, travaillée de cette maniére, prend un trés
beau poli, devient translucide dans les parties amincies, et sa
couleur varie du rouge-grenat au jaune-brun.
Le Rhinocéros blane est trés probablement Tl Unicorne ou
Licorne des anciens. Ctésias (410 av. J.C.) nous apprend que,
dés cette époque, on creusait dans la corne de Rhinocéros des
coupes qui avaient la reputation de mettre ceux qui s’en servaient
pour boire a labri de Veffet des poisons. C’est seulement au
moyen-dge que la défense de Licorne de mer ou Narwal (Monodon
monoceros) fut considérée comme ayant la meme propriéte,
et placée sur le front de la Licorne héraldique qui figure comme
support dans les armes de la Grande-Bretagne.
Mais l’ivoire de cette défense n’a jamais pu étre creusé en
forme de coupe a boire. Au contraire, les grandes cornes du
Rhinoceros simus, qui atteignent quelquefois, chez la femelle,
la longueur de 1™ 57, ont da frapper Pimagination des anciens,
et leur large base était tres propre a étre fagonnée en forme de
coupe.
EXPLICATION DES PLANCHES XXIX.-XXXI._
PLANCHE XXIX.
Rhinoceros simus cottoni, 6 et 2, du Bahr-el-Gazal, @aprés une aquarelle
de M. Terrier.
PLANCHE XXX.
Rhinoceros simus cottoni, 3, de profil, photographie prise au Bahr-el-Gazal.
PLraANcHE XX XI.
Rhinoceros simus cottoni, 8, le méme, vu de face, 11.
* Mohammed-al-Tounisi, ‘ Voyage au Wadai,” trad. francaise, 1851.
ConTENTS (continued).
February 2, 1909.
Mr. C, Tate-Regan, M.A., F.Z.S. Exhibition of two species of Char........++....+++05.
Mr. R. E. Holding. Exhibition of, and remarks upon, skulls and photographs of the
St. Kilda Four-horned Sheep ............... sg brat Sh apolar oe eek aeneee oieeas
Mr. Malcolm Maclaren, Account of a fight between a Whale and a Sword-fish ........ oe
Dr. C. W. Andrews, F.R.S., F.Z.S. Account of his visit to Christmas Island in 1908 ....
1, Preliminary Account of the Life-history of the Leaf-Insect, Phylliwm crurifolium Serville.
By H. S. Luten, Honorary Research Fellow in the University of Manchester.
(Gertie PXONGVPINUNT! i erste seveereershayerars iaie) dociic-s sie: sig cca gies el ete .
'
2. The Mammals of Matabeleland. By EH. C. Cuvss, F.Z.S., Acting Curator of the
AUlro desta Vs Currie sesuskeleysiersteraiere’e' <5 Secs: a2 cralie\o soe a,c uie,ra) 41a vebelee cites Rete NaN clea te Pyaatenes a0
ee ee ee ce ts ae ee ee ee oe oe oo
3. Report on Deaths which occurred in the Zoological Gardens during 1908. By H. G.
Primer, F.L,S.,'F.Z.8., Pathologist to the Society ..........ssecccevecnccsecsses
February 16, 1909.
The Secretary. Report on Additions to the Society’s Menagerie during the month of
dietneny MOS) osg eebatecac 3a aA Re LMEL AE Pie E Mic hy cil Geo. Psa las eree ae
Mr. C. Tate Regan, M.A., F.Z.S. Exhibition of sketches of, and remarks upon colour-
changes in some Fishes ...... aia cleie\s)(e\e i hetetal fete, chelelle)ot-tol/ojeleleel=teneteh tel steht teeta teteas oe
Mr. E. G. B. Meade-Waldo, F.Z.S. Remarks upon a letter from Dr. Einar Lonnberg
on the hunting of the Sea-Hlephant in South Georgia ........0++2 see cece cesses .
1, The Fauna of the Cocos-Keeling Atoll, collected by F. Wood Jones. By F. Woop Jonus,
B.Se., F.Z.S8., with the assistance of other Authors ............-- aranefabstesacetenctera ker ete
iw)
. Contributions to the Anatomy of certain Ungulata, including Zapirus, Hyrax, and
Antilocapra, By Frank E. Bepparp, M.A., F.R.S., F.Z.8., Prosector to the Society ..
8. Le Rhinocéros Blane du Soudan (Rhinoceros simus cottoni). Parle Dr. E, L, Trovessart,
C.M.Z.S., Professeur au Muséum d'Histoire Naturelle de Paris, (Plates XXIX.-XXXI.)
100
15
130
130
160
198
LIST OF. PiLAawiE
1909, pp. 1-200.
Plate Page
1, Drypanosomes: of Meleamdebercla sy. a /cciive ea. «estes cetera
II. Trypanosomes of Pike, Tench, and Bream. Trypanoplasms
Oy a el Menara coOIns Hit uateoM ooo dodo qaboee Ca icequaoe
III. Trypanoplasms of Tench, Bream, and Rudd........-..... 4 2
1V. Trypanoplasms of Pike, Tench, and Bream. Trypanosomes
OF Mere la 7.0358 ase tapecate love hn ato oleae uae occ paneas ene ene aod
(Vee Ey panosomes: of Melted Pench serucrerantertelces tees eeeteree
Vil SDianiomus galepordes GC sO ms SAEs w.1ciraldeneieiee 1nd eee \
Nie SDiapromus mints) Cha ON SAls. ac vielen aio tee eee Gad
VILL. Diaptomus stuhimanni Mrazek and D. simplex a. O. Sars .
LX Diaptomus cummngtoni G. O.Sarscere s-is s oes ote en
xX, -_Schizoperd (nopinatas Gr Oi SAGs sacrlese<toisieltsi ss «ici ae eee
Xl. Schizopera validior G. O. Sars, 8. consimilis G. O. Sars, and
SUNG ULOtG rs IOS Sansa & salantale ets arc) vk aa stk Seen ee
XII. Schizopera minuticornis G. O. Sars, 8. spinulosa G. O. Sars,
and 8, Jimbriata G. ONS ars So Ne oneness eee ee
XIII. Schizopera scalaris G. O. Sars and Ilyophilus perplexus
GO SS ang suis catieeseoierels ea sveusveve ave te cmmeiicee een eee
XIV. Cyclops leuckarti Ou C. emini Mrazek, and C. plete
Ors Op Samra g's Sra e On peels ete anes s ohare eal et eis es ernie rere ge |
XV. Cyclops tenellus G. O. Sars and C, albidus (Jurine) ...... + 31
XVI. Cyclops attenuatus G. O. Sars, C. varicans G. O. Sars, and |
Crexiguis G /OMSAES 9805 arms eicteseo ate shal oh ier es ie re
XVII. Cyclops cunningtont G. O. Sarsand C. pachycomus G. O. Sars
DAB Ne enalevam cies Ene OL IHS Go coos odhoosea be acenarac
XIX. Cyclops levimargo G. O. Sars, C. angustus G. O. Sars, and |
Caranispuiis: Gi Os Sars ieee aera cece ae eee tee
XX. Cyclops agiloides G. O. Sars, C. euacanthus G. O. Sars, and
Cicitnanas: Gh: O. ‘Sars ose eke tn acto a ee ee ea
XXI. Cyclops oligarthrus G. O. Sars, C. compactus G. O. Sars, and |
Craauis G-Os:Sars’ atm Sane 5 ooka y scien ene
XXII. LErgasiloides megacheir G.O. Sars .....
XXII. Hrgasiloides macrodactylus G. O. Sars and EE. ‘brevimanus
Ge CORB RES Ps: sis oie Sod ace este santtoegats is ace Aeane epee a ee eo )
XXIV. Gonadial Grooves of Aurelia aurita ........-00es.2+02.. 78
XXY.
KOGV Le UE NG QUECIOUIS io: crtaloperele: hive che aks wee eet eA Thee 91
XXVIII.
OQ OE RE EOL CUA OLUIDY Yo 6 Sedu Go oortns Aue sous boavon oo 103
XXSERG We eno cans) SIGNUS) COLLOMI ss) 4 Oe exntetes on slat sarcomere
See } Bhinoceros simus COLLOME, Sicha silent 6; Norarcter ait te 2 Soot Sapna se Ste {198
NOTICE.
The ‘ Proceedings’ for the year are issued in fowr parts, paged consecutively,
so that the complete reference is row P. Z. 8. 1909, p... . The Distribution
is as follows:— . ,
Papers read in January and February, issued in June.
a » March and April, » 9, August.
2 s, May and June, » » October,
a4 » November and December,,, ,, April.
‘ Proceedings,’ 1908, pp. 783-983, were published on April 8th, 1909.
The Abstracts of the papers read at the Scientific Meetings in
January and February are contained in this Part.
PROCHEDINGS
OF THE
GENERAL MEETINGS FOR SCIENTIFIC BUSINESS
OF THE
ZOOLOGICAL SOCIETY
OF LONDON.
1909.
Pages 201-544,
Part II. conTAINING PAPERS READ IN
MARCH anv APRIL.
AUGUST 1909.
PRINTED FOR THE SOCIETY,
SOLD AT THEIR HOUSE IN HANOVER SQUARE.
LONDON:
MESSRS. LONGMANS, GREEN, AND CO,
PATERNOSTER ROW.
| ae [Price Twelve Shilianys.]
LIST OF: CON EENTS:
1909, pp. 201-544.
March 2, 1909.
: Page
Mr. R. H. Burne, F.Z.8. Exhibition of, and remarks upon, certain elastic mechanisms in
Fishes and Reptiles... 0.2.0 cece cece cece ee ee ee eee ete eee cee cee e esse tee cees 201
Dr. R. F. Scharff, B.Se., F.Z.S. Exhibition of some long-bones and antlers of Reindeer
from County Cork.......- Beh ol a eas an Any cinmormo oot ee op cmc Cobo 76
Mr. R. I. Pocock, F.L.S., F.Z.S. On the Skulls of Leopards ...............+.. Sanus 204
1. The Development of the Subdivisions of the Pleuro-peritoneal Cavity in Birds. By
IMIAT GARB TER O GUN: cect rote Sancta meee aecie erclinvaliede ls stolter assis oMoletiteltis eto iegs ses eee 10
a ee eee ee ee eS e ee e
2. The Growth of the Shell of Patella vulgata L. By B.S. Russrut, M.A. (Plate XXXII.) 235 ;
3. The Life-History of the Agrionid Dragonfly. By Frank Baurour-Browng, M.A. (Oxon.),
F.R.S.E., F.Z.S. (Plates XXXITL. & XXIV.) -. eee eee ee cee cece rete ee cee 253
4. Growth-Stages in the British Species of the Coral Genus Parasmilia. By W. D. Lane,
M.A., F.Z.8., British Museum (Nat. Hist.). 2... cece eee eect ee cece ce eee eee e eens 285
March 16, 1909.
The Secretary. Report on the Additions to the Society’s Menagerie during the month of
February 1909 0.2. cs es ecine ce oe en sie seine we mien ase wievle «miele sein oilers 308
My. E. C. Chubb, F.Z.8. Exhibition of the skins and skulls of two fcetal Lions ....- eve oe) SOUS
The Secretary. Exhibition of a photograph of a young American Tapir ......-.++.++-+- 308
The Secretary. Note on a paper by C. Onelli in the ‘Revista del Jardin Zoologico de
HSTEMOS ANRC) faba ts hale aapanehers (eters site hate gave iePaee MEA NA obircremman tS oo Soe 30
The Secretary. Exbibition of a photograph of a small herd of Mountain Zebras ....-... 308
Vy p grap
The Secretary. Exhibition of a photograph of a female Giraffe captured in West Soudan.. 309 —
Dr. F. A. Bather, M.A., F.Z.S. Exhibition of a fossil Echinoid, Seudellina patella......-- 309
1. The Eetoparasites of the Red Grouse (Lagopus scoticus). By A. E. Sureuey, M.A.,
Hon.D.Sc., F.R.S., F.Z.8S., Fellow aud Tutor of Christ’s College, Cambridge, and
Reader in Zoology in the University. (Plates XXXV.—XALVIL.) ....-.0es- eee eees 309°
2, The Thread-Worms (Nematoda) of the Red Grouse (Lagopus scoticus). By A. BK. Suir.ey,
M.A., Hon.D.Se., F.R.S., F.Z.S., Fellow and Tutor of Christ’s College, Cambridge,
and Reader in Zoology in the University. (Plates XLVIIL—LV.).......+...-++++-5 335
3. The Tape-Worms (Cestoda) of the Red Grouse (Lagopus scoticus). By A. E. Surrey,
M.A., Hon.D.8c., E.R.S., F.Z.8., Fellow and Tutor of Christ’s College, Cambridge,
and Reader in Zoology in the University. With a Note by Wm. Bycravs, M.A.
(Plates sa V i Ske) eee eee ecieeenae I MAEM NA Rates Rea ea dod sg. oud s08 oc 351-
_ Internal Parasites of Birds allied to the Grouse. By A. HE. Smipiey, M.A., Hon.D.Se.,
E.R.S., F.Z.8., Fellow and Tutor of Christ’s College, Cambridge, and Reader in Zuology
ih the University. 21. sls mc'ele ows sie’ cle lei clei oo) wees ane olen) sient m= = nerfs nieel Ss ob
_ Ou a Fossil Bird from the Lower Pliocene. By W. P. Pycrarz, F.Z.S., ALLS. ..--..-. J68
+f
(Sp
6. On a Collection of Mammals from Western Java presented to the National Museum by
Mr. W. E. Balston. By Oxprimtp Tuomas, F.R.S., F.Z.S., and R. C. Wroventoy,
TOADS ieee ot alae Ft ae ete eee Sem OO art ad GOnraG Gorda“ dno < Bye
Contents continued on page 3 of Wrapper
1909.] | ON ELASTIC MECHANISMS IN FISHES AND A SNAKE. 201
March 2, 1909.
G. A. BouLencer, Esq., F.R.S., Vice-President,
in the Chair.
Mr. R. H. Burne, F.Z.S., exhibited and remarked on the followin g
specimens of certain elastic mechanisms in Fishes and Reptiles :—
“J. The ligamentum longitudinale ventrale of a Shad (Clupea
alosa).
The ligament lies in the cavity of the aorta suspended by a
longitudinal fold of its dorsal wall. In front it is firmly attached
to the basioccipital ; in the trunk region it is free except for the
mesentery-like fold of the aortic wall by which it is suspended
(text-fig. 20, M., p. 202); and in the tail region it becomes more and
more closely attached to the ventral surface of the vertebral
column, projecting merely as a ridge into the cavity of the aorta.
The ligament itself is a round or oval band of elastic tissue
forming in the trunk region a thickened edge to a longitudinal
curtain that divides the cavity of the aorta into two lateral
channels.
The ligament is extremely tight, so much so that when the
trunk muscles are removed, it draws the vertebral column into a
series of lateral serpentine curves. Its strength is also shown by
the large size of the elastic ligament between the neural spines.
In the trunk region the aorta is peculiarly spacious, filling a
wide channel upon the ventral surface of the vertebral column,
bounded by the flat lower surface of the vertebral column above
and by a series of quadrangular heemal arches at the sides and
below.
There is little doubt that this ligament is concerned in the
propulsion of blood within the aorta. ‘The following explanation
of its action was suggested to me several years ago by the late
Professor Charles Stewart.
When a Fish swims, lateral flexions, alternate from side to side,
pass regularly down the body from head to tail. This is most
manifest in such Fishes as the Eels and Dogfish, but in com-
paratively stiff fishes such as the Herrings the action is the same
although the chief and most powerful flexion takes place when
the wave of contraction nears the tail.
It is clear that the aorta will take part in these flexions—a
swing to either side sweeping in regular sequence from head to
tail (text-fig. 21, p. 203).
But the ventral ligament owing to its tension will remain
practically stationary and will in effect form a series of diagonal
curtains passing regularly in succession down the length of the
aorta. Hach curtain will of necessity push the blood before it.
The faster the fish swims the faster will become the blood
stream in the aorta.
This ventral longitudinal ligament is found in, I believe, all
Proc. Zoou. Soc.—1909, No. XIV. 14
202 MR. R. H. BURNE ON ELASTIC [ Mar. 2,
Herrings and also in the Sturgeons, and it is_ particularly
interesting to note that in both these groups of Fishes the aorta
has a similar character, being a wide capacious vessel enclosed
Text-fig. 20.
A transverse section through the vertebral column and aorta of a Shad,
in the region of the trunk.
A. Aorta. D.L. Dorsal elastic ligament. H.A. Hemal arch. M.* Suspensory
membrane of the ventral ligament. N.SP. Neural spines. V.B. Body of
vertebra. V.L. Ventral ligament.
within a more or less complete skeletogenous and unyielding
sheath. This is particularly so in the Sturgeon, where the vessel
1909. } MECHANISMS IN FISHES AND A SNAKE. 203
forms a closely adherent lining to a continuous cartilaginous
canal.
In any case the aorta is to a great extent cut off from the
direct effects of those internal movements and pressures to which
in Fishes the arterial circulation is probably to a large extent due,
as in the veins of Mammals.
We thus see in the case of these particular fishes the advantage
of some special mechanism to aid the circulation of the blood.
Text-fig. 21.
WANE
Diagrams of one and the same portion of the aorta, showing the relative positions
of the ventral ligament and aortic cavity during the lateral flexions of the body
in swimming.
The ligamentum longitudinale is apparently formed around,
though not actually from, the subchordal rod *—a structure that
occurs as a transitory foetal organ in Fishes and Amphibia, and
from its mode of development appears to be the vestige of the
epipharyngeal groove of Amphioxus.
* Franz, Morph. Jakrb. Bd. 25, 1897.
14%
204 MR. R. I. POCOCK ON THE [ Mar. 2,
2. The framework, muscles, and ligaments of a gill-pouch of
Raja maculata.
The expansion of the gill-pouch during the inspiratory move-
ment is due to a series of elastic ligaments acting upon the
branchial rays, drawing them away from the central member of
the series.
3. An elastic modification of the anterior wall of the capsule of
the costo-vertebral joints in a Python (P. sede).
This part of the capsule-wall is greatly thickened and consists
almost entirely of elastic fibres arranged parallel to one another
and to the long axis of the rib.
The action of this plate of elastic tissue would no doubt be to
protract the rib upon the relaxation of its retractor muscles.
In Snakes forward locomotion is largely due to the forcible
retraction of the ribs, their protraction being rather in the nature
of a recovery to a state of repose—a movement requiring com-
paratively little force.”
Dr. R. F. Scharff, B.Sc., F.Z.S., exhibited some long-bones and
antlers of Reindeer from a cave in County Cork in Ireland.
He stated that Reindeer bones had been discovered sparingly
in caves in the Counties of Sligo and Clare along with those of
the Greenland Lemming (Dicrostonyx torquatus). During the
Co. Cork cave explorations, the results of which will shortly be
published in detail, Reindeer remains were found in extreme
abundance. They were associated with the bones of Hyena and
two species of Lemming.
The main object of exhibiting the Reindeer bones was to draw
attention to the fact that many of them showed distinct traces
of having been gnawed by some other animal. Dr. Scharff had
suggested that the tooth-marks might have been produced by
young hyenas, but was disposed to accept the view that they
were produced by Rodents. The marks on the antlers seemed
to differ from those on the long-bones, the latter having often
been gnawed through entirely in the middle.
Mr. Pocock on the Skulls of Leopards.
Mr. R. I. Pocock, F.L.S., F.Z.S., the Superintendent of the
Gardens, exhibited the skulls of some West African Leopards
which had come from Cette Cama in the Gaboon, and made the
following remarks :—‘“ I owe to Mr. Edward Gerrard, taxidermist,
of Camden Town, the opportunity of examining these skulls,
which he sent to me on account of their unusual size. He was
struck by the fact that three of them surpass in dimensions all
the Leopards’ skulls out of the very large number that have passed
through his hands, and are particularly remarkable for the length
and thickness of the canine teeth. A comparison between their
measurements and those cited in Rowland Ward’s ‘ Records of
1909. ] SKULLS OF LEOPARLS. 205:
Big Game,’ p. 500, 1907, shows that all three exceed both in
length and width the largest therein recorded, the measurements.
of which were taken from an example from the North-Western
Provinces of India. They also surpass in dimensions all the
Text-fig. 22.
Skull of Leopard (Felis pardus leopardus) from Cette Cama. Ventral view.
(No. 1 in table on page 208.)
Leopards’ skulls I have seen in the collection of the British
Museum and all those in the collection of the Society. More-
over, although their shape indicates very considerable develop-
ment of the temporal muscles, the persistence of the divisional
lines between the bones of the brain-case and the unworn character
206 MR. R. I. POCOCK ON THE [ Mar. 2,
of the teeth show that they are not skulls of old Leopards, and
suggest that they might have grown even larger if the animals
had not been killed. In one indeed (No. 3) the occipital suture
is still open. The locality proves practically beyond doubt that
they belong to the tropical West African race of the Leopard
(felis pardus) which should be called /. pardus leopardus*.
Text-fig. 23.
Skull of Leopard (Felis pardus pardus) trom Mhow, C.P., India. Ventral view.
(No. 9 in table on page 208.)
From the largest Leopard’s skull in the collection of the Society,
they exhibit some very striking differences as the subjoined table
* Pocock, P. Z. S. 1907, p. 675.
La
1909. | SKULLS OF LEOPARDS. 207
of measurements shows (p. 208). This skull itself nearly reaches
record dimensions since it exceeds in zygomatic width by half an
inch the largest recorded by Rowland Ward (71 : 68) and falls only
a quarter of an inch short of it in total length (10:103). It
belonged to an exceptionally fine male from Mhow in the Central
Provinces of India, presented to the Society by Mr. Pelham, This
Leopard was of the typical race /’. pardus pardus. Not only is
this skull from Mhow broader across the premaxille than the Cette
Cama specimens, but it also differs from them in having much
smaller auditory bulle, the width of these being less than the width
of the basioccipital area between them, instead of being greater ;
but most of all does it differ in the width and shape of the meso-
pterygoid fossa, which is wider in front than behind and has a
sinuous concavo-convex edge owing to the incurvature of the
pterygoid bones; whereas this fossa in the Cette Cama skulls has
almost parallel sides and is comparatively long and narrow though
varying individually in these respects. It is also of interest to record
that a Leopard’s skull of average dimensions belonging to a male,
presented to the Society by Mr. F. H. Melland and captured on the
Luangwe River in North-east Rhodesia, has the mesopterygoid
fossa shaped more like that of the specimen from Mhow than like
the fossa of those from Cette Cama; and that in the equality in
width between the auditory bull and the intervening area of the
basioccipital, it occupies an intermediate stage. As I have already
stated *, this Luangwe Leopard also differs from the typical West
African form in colour, its yellow hue recalling that of most
Indian specimens, although the spots are much smaller. This fact
and the differences in the skull above recorded indicate that it
represents a distinct subspecies from /”. pardus leopardus ; but in
the present very unsatisfactory state of our knowledge of East
and South African Leopards, I do not feel justified in applying
to 1t a name, either new or old.
Mr. Gerrard also sent to me with the three above mentioned
skulls from Cette Cama, doubtless belonging to male specimens,
a single skull of what I take to be a female animal, which was
shipped from the same port at the same time together with some
spoils of Gorillas and Chimpanzees. The closure of the cranial
sutures shows that the specimen was adult and yet it is re-
markably small, being 3 inches shorter and more than 2 inches
narrower than the smallest of the three large ones (cf. measure-
ments). This skull lends support to the oft-repeated statements
of sportsmen that two kinds of Leopards, larger ones called
Panthers and smaller ones called Leopards, occur in the same
localities.
So far as size goes this small specimen approaches much nearer
the typical skull of the dwarf Leopard of Somaliland described by
Thomas as F, pardus nanopardus, a Leopard which, together with
* P. Z. S. 1907, p. 676.
[ Mar. 2,
MR. R. I. POCOCK ON THE
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1909. ] SKULLS OF LEOPARDS. 209
its normal-sized congener coming from Somaliland, furnishes the
most striking instance known of a larger and smaller form
occurring in the same country.
In the subjoined table I have placed side by side for comparison
measurements in English inches* of eight skulls of African
Leopards and of one Indian example, namely, the above mentioned
one from Mhow. Nos. 1-4 are the four examples from Cette
Cama lent to me by Mr. Gerrard. No.5 is Mr. Melland’s Luangwe
River small-spotted specimen. No.6 is an example in the British
Museum sent from the French Congo by Mr. G. L. Bates.
Although smaller than the large Cette Cama skulls, this presents
the same general features and has the mesopter yeoid fossa narrow,
with nearly parallel edges. No. 7 isa specimen from Fort Man-
ning in Nyasaland, in yee British Museum. Although considerably
smaller than No. 3 of the Cette Cama series and larger than the
Luangwe River example, this skull has the mesopterygoid fossa
of the same actual width. The skull of another Leopard from
Zomba in Nyasaland, however, has the mesopterygoid fossa broad,
namely + of an inch, that is to say very nearly as broad as in the
considerably larger iiactiewn example from Mhow. It will be seen
therefore that the marked distinction in width and shape of the
mesopterygoid fossa between Indian and West African Leopards
breaks down in East and South African specimens, as is attested
by the skull from Zomba, the one from Fort Manning, and also
one from Cape Colony in the British Museum. The fossa in
question is wide in all the Asiatic Leopards’ skulls that I have
seen, including specimens from India, North China (7. pardus
fontanieri), Hong Kong, and of the Black variety from Malacca.
It is also broad in the skulls of Jaguars (/. onca) and points to
closer relationship, also borne out by the pattern, between Jaguars
and Asiatic Leopards than between Jaguars and West African
Leopards, a conclusion to be expected from geographical dis-
tribution. On the other hand, the anterior border of the fossa is
at all events as a rule produced backwards into a sharply pointed
angle in Asiatic Leopards, whereas in African Leopards it is
generally at least markedly straighter, sometimes indeed slightly
notched, and in Jaguars this edge is usually notched, as it is also
in Lions.
In conclusion it may be said that Asiatic Leopards (/. p. pardus,
fontunieri, etc.) have the mesopterygoid fossa wide, with arcuate
lateral edges and the anterior edge strongly angled. West African
Leopards (fF. p. leopardus) have the mesopterygoid fossa narrower,
with subparallel lateral edges and a straighter anterior border,
whereas in Hast and South African animals some present one
form of fossa and some the other; but I have not seen sufficient
material to contribute anything further to the elucidation of the
local races of Leopards, occurring between Abyssinia and Cape
Colony.
* An English inch=25 mm.
210 MISS M. POOLE ON THE DEVELOPMENT OF THE __ [ Mar. 2,
The following papers were read :—
1. The Development of the Subdivisions of the Pleuro-
peritoneal Cavity in Birds. By Marcarer Poois.*
[Received February 9, 1909.]
(Text-figures 24-37.)
The subject of this paper was suggested to me by Professor
Bourne, and it was by his advice that I undertook a reinvestiga-
tion of the development of the subdivisions of the body-cavity in
birds; my object being to confirm and enlarge the results put
forward in a paper on the subject by Mr. G. W. Butler in these
‘Proceedings.’ At that time I knew of no other work on this part
of the question; and it was only when IJ had almost completed my
study of the development of these structures, that Mr. Beddard
kindly referred me to the work of Dr. Bertelli. This I found was
published in an Italian journal not—so far as I could discover—
taken in by any of the scientific libraries either in Oxford or
London ; and it was only by the very great kindness of Dr. Bertelli,
who sent me a copy, that I was at last able to seeit. I then
found that he had worked out and described in great detail and
with admirable clearness, almost the whole of the development
upon which I was engaged. But since this valuable paper is
practically inaccessible to English students, Dr. Bourne advised
me to publish the results of my own investigations, which I had
arrived at independently. I may here state that my results
entirely confirm those of Mr. Butler, and, except in two small
points, those of Dr. Bertelli also.
;: The work was done in the Zoological Laboratory at Oxford py
the kindness of Professor Bourne, and I wish to take this oppor-
tunity of expressing my warmest thanks to all in the Department
—and especially to Mr. Goodrich—who have helped me by
continual suggestions and advice.
I wish likewise to express my gratitude to Dr. Bertelli for his
kindness, not only in sending me a copy of his work, but also for
his permission to make any use of his observations in my own
paper.
NOMENCLATURE.
The pulmonary aponeurosis of Huxley [12] and Butler [9]—
diaphragme pulmonaire of Sappey [16]—Bertelli [6, 8] calls the
diatramma ornitico.
The oblique septum of Huxley, employed by Butler, Beddard and
others—the diaphragme thoraco-abdominal of Sappey—Bertelli
does not consider to be a septum at all, but only the ventral walls
of the intermediate air-sacs together with the posterior wall of
* Communicated by Prof. G. C. Bourne, D.Sc., F.Z.S.
1909. ] SUBDIVISIONS OF THE BODY-CAVITY IN BIRDS. 211
the pericardium, Butler takes the pulmonary aponeurosis and
the oblique septum as forming together a single septum, which he
terms the avian diaphragm. The backward prolongation of this
in connection with the development of the posterior air-sacs, he
calls the oblique abdominal septum. Since, however, Bertelli’s
diaframma ornitico is but half of Butler’s avian diaphragm, I shall
use neither of these terms, but call the whole of Butler’s avian
diaphragm together with his oblique abdominal septum, the post-
pulmonary septum ; this, in the early stages of its development,
corresponds to what Bertelli calls the “ diaframma primario.”
For the air-sacs I shall use the terms employed by Bertelli in
his earlier paper (7]:—Anterior intermediate, posterior inter-
mediate, and posterior; except for the latter, which he calls
abdominal, Butler uses the same names.
The blind anterior diverticula of the peritoneal cavity I shall
call, after Butler, the pulmo-hepatic recesses; these are the
“‘ yecessi polmonali-enterici ” of Bertelli. Butler’s pulmo-hepatic
ligaments correspond to Bertelli’s ventral pulmonary ligaments,
and I shall use these latter terms in describing the development.
Finally, I shall call, with Butler, the horizontal septum of
Beddard [1] the post-hepatic septum; since this term, while
indicating its essential position, does not depend upon its direction
which is very variable.
I. Empryowocy.
The Post-pulmonary Septum.
As it is important to follow the growth of the lungs from their
first appearance, Bertelli begins his observations on chicks of 60
hours’ incubation, though the septum does not begin to develop
until much later. At this stage the bronchi are not yet
formed, but the mesoderm into which they will grow appears
as slight folds (meso-laterali) on the lateral faces of the median
dorsal mesentery (setto mesenterico), projecting into the cceelomic
cavity. That on the right side is united ventrally with the
septum transversum, this being the vertical partition posterior
to the pericardium which divides the latter from the peritoneal
cavity, and in its more dorsal portion carries the two ductus
Cuvieri inwards to the heart. After three days’ incubation, the
folds have increased in size, and both are now united ventrally
with the septum transversum. The bronchi are formed, and
have penetrated into the mesodermal folds, and so divided the
latter each into three regions: a posterior, dorsal pulmonary
ligament, a median, pulmonary rudiment (abbozzo polmonale),
and an anterior, ventral pulmonary ligament. Between the
ventral pulmonary ligaments on each side, and the median dorsal
mesentery, lie the narrowed anterior prolongations of the peri-
toneal cavity—the pulmo-hepatic recesses. On the fourth day, the
relations between these parts remain the same, but in a chick of
five days’ incubation the pulmonary rudiments have greatly
212 MISS M, POOLE ON THE DEVELOPMENT OF THE [ Mar. 2,
increased in size, and the ventral pulmonary ligaments have
united ventrally with the lower edge of the median dorsal mes-
entery, the dorsal surface of the liver, and the pericardio-pleuro-
peritoneal membrane. The latter is what is called in younger
stages the septum transversum ; it now forms a mass of tissue
separating the pericardium from the pleural and the peritoneal
regions of the celom. Posteriorly, the ventral pulmonary
ligaments spread out laterally to unite with the antero-lateral
corners of the now well-developed liver; and at the same time the
dorsal pulmonary ligaments and the lung rudiments themselves
assume an almost transversely horizontal position; so that in this
region there is now a septum across the ccelomic cavity, complete
except at its extreme lateral margins where the pleural and
peritoneal divisions communicate by a narrow aperture on either
side. Ata later stage these passages will be closed by the union
of the ventral pulmonary ligaments and the lateral body-walls,
and thus the post-pulmonary septum—bBertelli’s ‘“ diaframma
primario ’—will be formed.
The pronephric folds appear at this stage as thickenings of the
connective tissue on the external walls of the anterior part of the
pleuro-peritoneal cavity; posteriorly they are seen in close con-
nection with the developing Wolffian bodies on the dorsal wall of
the ccelomic cavity.
Text-fig. 24 is of a transverse section of a chick of 5 days’
Text-fig. 24.
Transverse section of a chick of 5 days’ incubation, through the anterior region
of the liver; anterior face.
For explanation of abbreviations in text-figures see p. 235.
incubation, taken through the region where the septum will be
formed, the anterior face being shown. The developing pro-
nephros is seen on the dorsal wall of the pleuro-peritoneal cavity.
1909. ] SUBDIVISIONS OF THE BODY-CAVITY IN BIRDS. 213
Text-fig. 25 isa slightly diagrammatic reconstruction from longi-
tudinal horizontal sections of a chick of six days’ incubation, seen
from the ventral side. The pericardium and heart, the ventral
part of the alimentary canal, and all the liver except the most
dorsal corner of the right lobe have been removed.
Text-fig. 25.
\
Sp.rnl.
Slightly diagrammatic reconstruction from horizontal longitudinal sections of a
chick of 6 days’ incubation ; seen from the ventral side. The pericardium
and heart, the ventral part of the alimentary canal, and all the liver except
the most dorsal corner of the right lobe have been removed.
At this stage the two pairs of intermediate air-sacs first make
their appearance, lying —together with the posterior air-sacs which
are developed even earlier—in the ventral pulmonary ligaments.
The pronephric folds are now seen, supporting the Miillerian ducts
214 MISS M, POOLE ON THE DEVELOPMENT OF THE [ Mar, 2,
Text-fig. 26.
\ ,
Wd 0 } io <
Pppm. Pn.rs. Ht. ‘Es. “st >
Transverse section of a chick of 7 days’ incubation, through the posterior part of the
pericardium and anterior part of the liver; anterior face.
Text-fig. 27.
Longitudinal vertical section of a chick of 7 days’ incubation, taken a little to the
left of the middle line; seen from the right side.
1909. | SUBDIVISIONS OF THE BODY-CAVITY IN BIRDS. 215
at their extremities, projecting from the dorsal walls of the pleural
parts of the coelomic cavity. Posteriorly they unite with the
pericardio-pleuro-peritoneal membrane, and thus for a short space,
the pleural and peritoneal cavities are separated from one
another, ‘This is shown in text-fig. 26, which is of a transverse
section of a chick of 7 days’ incubation. On the left side of
the section this connection has disappeared ; on the right side it
is lost a few sections further back, and then the two cavities
communicate freely. Here the pronephric folds hang free
from the lateral body-walls into the ccelom—as shewn on the
right side of the section—until they unite with the Wolffian
bodies.
Text-fig. 27 is of a vertical longitudinal section through a
chick of the same age.
On the eighth day of incubation, immediately above their
connection with the pericardio-pleuro-peritoneal membrane, the
ventral pulmonary ligaments have united with the lateral body-
walls, and thus formed a part of the post-pulmonary septum.
The air-sacs are now quite large, and by their backward growth
have pushed the connective tissue of the more median portions of
the ventral pulmonary ligaments, together with the tissue of the
pericardio-pleuro-peritoneal membrane at the posterior edge of the
lungs—which at this time cannot be distinguished from the
pulmonary tissue itself—before them into the peritoneal cavity.
This post-pulmonary tissue on either side, containing the air-
sacs, forms the antero-lateral wall of the pulmo-hepatic recess.
Text-fig. 28, p. 216, is a reconstruction similar to text-fig. 25 of
a chick 8 days after the commencement of incubation. The pleural
cavities are seen closed ventro-posteriorly by the union of the
ventral pulmonary ligaments with the lateral body-walls. More
dorsally, however, the communications with the peritoneal cavity
are widely open, as is indicated by arrows.
Ventrally, the pronephric folds are attached to the dorsal
surface of the ventral pulmonary ligaments where these unite with
the lateral body-walls; behind this connection the folds hang
free into the celom, bearing the Miillerian ducts at their
extremities, until on the dorsal surface they unite with the
Wolttian bodies.
On the tenth day of incubation, the pleural and peritoneal
cavities are completely separated by the dorsal extension of the
union between the ventral pulmonary ligaments and the lateral
body-walls on either side. Besides this, however, the backwardly
growing post-pulmonary tissue containing the air-saecs, which
appeared on the eighth day as projecting freely into the peritoneal
cavity, has become united with the lateral body-walls behind
the ventral pulmonary ligament connection. As this union takes
place gradually during the ninth day, from before backwards, the
anterior prolongations of the peritoneal cavity—previously
forming the lateral boundaries of the post-pulmonary masses of
tissue—become completely obliterated. Thus the post-pulmonary
septum, Bertelli’s ‘‘ diaframma primario,” is established.
216 MISS M. POOLE ON THE DEVELOPMENT OF THE [ Mar. 2,
The anterior intermediate air-sacs are at this stage large oval
vesicles lying in the post-pulmonary septum néar the middle line
and close against the base of the lungs. The posterior inter-
mediate air-sacs are much smaller, and enclosed in the extreme
lateral portions of the same septum. The posterior air-sacs,
arising from the postero-lateral corners of the lungs, are very
large and project backwards for a considerable distance into the
peritoneal cavity. Bertelli says that their posterior ends project
Text-fig. 28.
moose
— noe
EEA
=
oe
Bia
Reconstruction, similar to text-fig. 25, of a chick of § days’ incubation.
beyond the diaphragmatic tissue and thus ‘‘resta libero nella
cavita addominale tra la parete laterale di questa, il corpo di
Wolff e Vintestino a sinistra, tra la parete laterale dell’ addome, il
corpo di Wolff e il fegato a destra.” On this point my own
observations differ slightly from his. I believe that the posterior
ends of these air-sacs never extend beyond the hindmost edge of
the post-pulmonary septum. However far these air-sacs extend
among the abdominal viscera, they always push before them a
layer of connective tissue covered with coelomic epithelium.
Text-fig. 29 is of a reconstruction, similar to text-figs. 25 and 28,
1909. ] SUBDIVISIONS OF THE BODY-CAVITY IN BIRDS. 217
of these structures in a chick of 10 days’ incubation. In addition,
this figure shows the origin of the first costo-pulmonary muscles
from the fifth ribs, which run into the post-pulmonary septum
following the ventral and posterior border of the lungs. These
will be supplemented by others arising later from the third,
fourth, and sixth ribs, and give rise to the aponeurotic mem-
brane, which in the adult covers the postero-ventral face of the
lungs and separates these organs from the intermediate air-sacs.
The pronephric folds are attached ventrally to the antero-
lateral walls of the post-pulmonary septum, and dorsally are con-
nected with the external margins of the Wolffian bodies.
Text-fig. 29.
Reconstruction, similar to text-figs. 25 and 28, of a chick of 10 days’ ineubation.
The post-pulmonary septum, now fully formed from the median
dorsal mesentery in the middle line, the pericardio-pleuro-
eritoneal membrane with the ventral pulmonary ligaments, and
slightly also from the pronephric folds laterally, constitutes a
complete partition dividing the celomic cavity into a pleural
and a peritoneal portion. JBertelli calls this septum the
Proc. Zoot. Soc.—1909, No. XV. 15
218 MISS M. POOLE ON THE DEVELOPMENT OF THE [ Mar. 2,
‘« diaframma primario ” because he says it isa temporary structure
which will later give rise to the “ diaframma definitivo” and also
to the ventral walls of the intermediate air-sacs. At this stage—
ten days after incubation—he says the anterior intermediate air-
sacs lie enclosed in the primary diaphragm, and by splitting it
into two layers, form with their dorsal walls the definitive
diaphragm and with their ventral walls the so-called “ diaframma
toraco-addominale.” The posterior intermediate and the posterior
air-sacs likewise split the primary diaphragm into two layers; and
behind this he describes them as lying in the lateral abdominal
walls, covered by their peritoneum, and bounded externally by the
connective tissue which surrounds the muscles of the body-wall.
Now I do not find any signs of these air-sacs actually penetrating
into the walls of the peritoneal cavity so as to lie between its
bounding celomic epithelium and the musculature of the body-
wall. According to my observations, the air-sacs always remain
enclosed in the backward prolongations of the post-pulmonary
septa, and this tissue becomes closely attached—throughout its
entire thickness—to the lateral body-walls. Between the post-
pulmonary tissue surrounding the air-sacs and the connective
tissue of the body-wall, in all my sections there is a darkly-
staining streak, showing—I believe—where the two layers of
ceelomic epithelium have fused together (text-figs. 29, 30, 31, cf).
And I differ a little from Bertelli as to the formation of his
“ diaframma definitivo” from the antero-dorsal portion of the
“ diaframma primario ” ; though this divergence of opinion depends
entirely upon my own conception of the essential nature of a
coelomic septum. I consider that sucha septum should be covered
on both surfaces by peritoneum, and really separate one division
of the celom from another; and therefore I prefer—following
Butler—to regard the whole post-pulmonary mass of tissue, with
the intermediate and posterior air-sacs enclosed between its
antero-dorsal and postero-ventral walls—the “ diaframma ornitico”
and the so-called *‘diaframma toraco-addominale” respectively of
Bertelli—as a single septum. Except for the presence of the costo-
pulmonary muscles in the antero-dorsal half of this septum,
there is nothing to distinguish it, as the true diaphragm, from
the postero-ventral portion, in the first stages when the splitting
into two layers occurs with the development of the air-sacs.
Besides which, the ‘“diaframma ornitico” of Bertelli only
separates the pleural cavities from the cavities of the inter-
mediate air-sacs, and is consequently covered only on one face
by peritoneal epithelium; the other surface being invested by
the endoderm of the air-sacs. What I have described as the
post-pulmonary septum, on the other hand, does really separate
the pleural from the peritoneal cavity, and is covered ventrally,
as well as dorsally—except where the septum is continuous with
the tissue of the lungs—by ceelomic epithelium. The interposition
of the air-sacs between its two surfaces, does not, I believe,
essentially alter the nature of the whole. My observations
1909. | SUBDIVISIONS OF THE BODY-CAVILY IN BIRDS. 219
entirely confirm Butler’s conclusion, that “the avian diaphragm
is seen to be completed as a single structure, and its separation
into its two lamine is a secondary detail arising in connection
with the development of the two pairs of intermediate air-sacs,
which first penetrate it at a time when no distinct line can be
drawn between the tissue that goes to form it, and that of the
developing lung itself.”
The adult arrangement of these structures is shown in text-fig. 32,
p. 222, which is of a section of a Rook through the region of the
liver and gizzard, seen from the ventral side. The post-pulmonary
tissue is seen united to the lateral body-walls for a very consider-
able distance on either side, and ventrally and towards the
middle line is pushed out irregularly into the peritoneal cavity by
the growth within it of the posterior air-sacs. The latter,
however, as I have already said, never project beyond the edge of
the septum in which they he, but always carry a layer of con-
nective tissue and cceelomic epithelium in front of them.
Text-fig. 30 is of a transverse section of a chick, 13 days after the
peginning of incubation, through that part of the post-pulmonary
Text-fig. 30.
D.ms.
1
'
1
I
t
1
Uy
U
Phurs:
Transverse section of a chick of 13 days’ incubation, through the region of the
pericardium and anterior intermediate air-sacs ; seen from behind.
septum which encloses the anterior intermediate air-sacs. The
lungs are seen now lying in completely closed pleural cavities,
separated from each other in the middle line by the median
dorsal mesentery.
iL
220 MISS M. POOLE ON THE DEVELOPMENT OF THE [ Mar. 2,
The post-pulmonary septum stretches across the coelomic cavity
from one side to the other, closely fused to the lateral body-walls
throughout its entire thickness. In it, near the middle line, lie
the anterior intermediate air-sacs. The costo-pulmonary muscles
pass inwards from the ribs dorsal to the air-sacs following the
ventral border of the lungs. They really form a continuous
layer dorsal to the cesophagus, but in the section figured they are
interrupted by the ostia of the anterior intermediate air-sacs.
Forming the ventral boundaries of the post-pulmonary septum
are, medianly, the pulmo-hepatic recesses, and laterally, the
abdominal cavity. Behind the pericardium at this stage the
posterior intermediate air-sacs, still comparatively small, lie in the
periphery of the septum, and more posteriorly the posterior air-
sacs occupy the same position. These however have extended
both ventrally and also towards the middle line, always carrying
some tissue of the post-pulmonary septum with them, and have
united with the median mesentery of the gizzard and liver, dorsal
to these organs and ventral to the Wolffian bodies. This union
forms the beginning of the post-hepatic septum to be described
below. Text-fig. 31 is of a transverse section of the same chick of
13 days’ incubation, taken through the region of the spleen,
and showing the inwardly expanded post-pulmonary septum—
containing the greatly enlarged posterior air-sacs—united to the
gastro-hepatic ligament.
The Post-hepatic Septun.
This septum is completed in the chick on the 15th or 16th day
of incubation. The first to be formed is the antero-dorsal moiety,
and this is fully developed by the 15th day. Text-fig. 31, as I
have already described, shows the formation of this part by the
inward expansion of the two halves of the post-pulmonary septum
(those parts termed by Butler the ‘oblique abdominal septa’)
towards the middle line, where they meet and fuse with the
mesentery of the gizzard and liver. This lateral extension is
due to the greatly increased size of the posterior air-sacs enclosed
within the septum.
Ventral to this connection the gizzard is seen in text-fig. 31
included within the median mesentery, which is therefore split
into two layers. Posteriorly this mesentery expands laterally
before it becomes attached to the ventral body-wall; and at a
later stage its lateral edges become fused to the side walls of the
peritoneal cavity, thus constituting the postero-ventral portion of
the post-hepatic septum. When fully formed, this is a membrane
passing obliquely from the antero-dorsal to the postero-ventral
walls of the peritoneal cavity and enclosing within its thickness the
gizzard in the more posterior portion. It is a complete partition
except on the left side where the dorsal and ventral components
never quite meet, and so there remains always a narrow com-
munication between the post-hepatic intestinal cavity and the
1909. ] SUBDIVISIONS OF THE BODY-CAVITY IN BIRDS. 221
pulmo-hepatic recess of that side. Text-fig. 33, p. 223, shows very
diagrammatically the divisions of the ccelom in the adult. bird in
longitudinal vertical section. Text-fig. 34 is of a chick 17 days
after the beginning of incubation, bisected longitudinally a little
to the left of the middle line, and seen from the right side. The
arrangement of the septa is the same as in the adult fowl but is
shown more clearly, since in the latter the mesenteries are often
loaded with fat. The gizzard is seen enclosed in the post-hepatic
septum just dorsal to the attachment of the latter to the ventral
body-wall, and in front of the umbilicus. Dorsally the septum is
seen formed first by that part of the median mesentery which
supports the spleen in the left pulmo-hepatic recess (see also
text-fig. 31), and then by the post-pulmonary septum which is
Text-fig. 31.
=.
SH
Li
NS Yj
Ye
(Nw
Transverse section of a chick of 13 days’ incubation, through the connection between
the posterior part of the post-pulmonary septum and the mesentery of the
liver and gizzard; seen from behind.
itself united dorsally with the vertebral column by means of the
median dorsal mesentery (see text-fig. 30, p. 219). By this time the
air-sacs have increased so much in size, as completely to obliterate
all the connective tissue of the post-pulmonary septum which
at first surrounded and separated them from each other; and
the walls of the adjacent air-sacs have become closely applied to
each other, As a result of this, the post-hepatic septum appears
in the figure to be completed dorsally by the posterior wall of the
posterior intermediate air-sac together with the fused anterior
222 MISS M. POOLE ON THE DEVELOPMENT OF THE [ Mar. 2,
wall of the posterior air-sac. From an examination of the earlier
stages, however, it is evident that the dorsal component of the
post-hepatic septum is really formed by the whole of the more
posterior portion of the post-pulmonary septum, with its contained
air-Sacs.
Text-figs. 32, 33, and 34 also show those peritoneal cavities,
called by Butler the ventral liver-sacs, in which the two liver-
Text-fig. 32.
tly
thon:
Adult Rook bisected horizontally through the region of the gizzard and liver-lobes,
and slightly dissected anteriorly to expose the lungs; seen from the ventral
side.
lobes are seen to lie upon opening a bird from the ventral side.
These are clearly formed by no special development of septa, but
are simply those parts of the peritoneal cavity which, by the ventral
attachment to the body-wall of the post-hepatic septum, become
shut off anteriorly between this septum and the posterior wall of
1909. ] SUBDIVISIONS OF THE BODY-CAYVITY IN BIRDS. 223
the pericardium. Their lateral walls are formed by the post-
pulmonary septum, and the median partition between them is the
ventral mesentery or falciform ligament.
Text-fig. 33.
Jeti (OF es spt.
Abd.c -7 fe
Ph. rs:
Bhp. sp ryy.s.
Diagrammatic plan showing the subdivisions of the celom in a bird in
longitudinal section.
Text-fig. 34.
Ppl.spe. . Ing.
va Puy. as. /
Chick of 17 days’ incubation bisected longitudinally a little to the left of the
middle line; seen trom the right side.
These figures show likewise that the pleural cavities persist
even in the adult; though they tend to get somewhat obscured
by the fusion here and there of the adjacent layers of pleural
epithelium, and the development of connective-tissue strands
across the cavities.
224 MISS M. POOLE ON THE DEVELOPMENT OF THE _ [ Mar. 2,
IJ. Aputr Anatomy.
The Post-pulmonary Septum.
Bertelli [8] gives an extremely clear description, accompanied
by an excellent drawing, of his “diaframma ornitico” in the
Fowl, with the attachments of all the costo-pulmonary muscles.
Since, however, Sappey [16] has given such an elaborate account
of the whole post-pulmonary septum in the Duck, and Huxley [12]
has done the same in Apteryx, I do not propose to go over the
same ground again here. But as—in spite of the recent embryo-
logical work on the subject—there appears to be still some
uncertainty as to the nature of the cavity within the post-
pulmonary septum (that is to say between the pulmonary apo-
neurosis and oblique septum of Huxley), it may be as well to
point out those parts of the descriptions given by Huxley and
others which embryology has shown to be incorrect.
Huxley [12] in describing very clearly the partition which
shuts off from the general visceral cavity a space enclosing the
lungs and either most or all of the air-sacs on each side, appar-
ently assumes, like Sappey, that this partition consists of two
distinct septa, between which lies a cavity which is a division of
the celom. For between the pulmonary aponeurosis dorsally
and laterally, and the oblique septum ventrally and posteriorly,
the lateral body-wall and the median dorsal septum, Huxley
describes a space which he calls the subpulmonary chamber.
This, he says, “is divided into four loculi by three dissepiments,
which pass transversely from the lateral face of the oblique
septum to the mesial face of the pulmonary aponeurosis. Each
loculus lodges one of the four postbronchial saccular diverticula
of the wall of the lung, constituting the proper air-sacs, which
thus fill up the subpulmonary chamber, between the insertion of
the bronchus and its posterior extremity. .... Thus, that part of
the thoraco-abdominal cavity which lies dorsad and anterior to
the oblique septum lodges no other viscera than the lungs and the
air-sacs, and may be distinguished as the respiratory cavity, from
the cardio-abdominal cavity which contains the heart and the rest
of the viscera, and lies below and behind the oblique septum.
The respiratory cavity is further divided into two lateral chambers
by the median dorsal septum; and each of these chambers is
subdivided by the pulmonary aponeurosis into two stories, of
which the upper is occupied by the lung, and the lower by the
loculi with their contained air-sacs.” In the adult bird these
structures do appear as above described, but from working out
the development, it is at.once clear that the cavity lying between
the pulmonary aponeurosis and the oblique septum on each side,
is not a division of the celom lined by peritoneum, but merely
the cavities of the three posterior air-sacs, and therefore lined by
endoderm. The dissepiments which are described as subdividing
the subpulmonary chamber into loculi, are only the walls of the
same air-sacs, which by the growth of the latter have become
1909. | SUBDIVISIONS OF THE BODY-CAVITY IN BIRDS. 225
closely applied to one another, and so have obliterated the con-
nective tissue of the post-pulmonary septum which at first separated
them. The pulmonary aponeurosis and the oblique septum, as
already shown, together constitute a single partition across the
celomic cavity, into the thickness of which the intermediate and
posterior air-sacs push their way as they develop; the latter are
never at any stage surrounded by a part of the body-cavity.
Roché [15] and, in the last year, Miiller [14] describe the sub-
divisions of the ccelom in the same way.
Beddard [5] describes two interesting variations ip the adult
arrangement of the oblique septa. In the Emu, for example, he
says, ‘‘ the posterior part of the oblique septum is free from the
abdominal walls, ending, in fact, in a free edge within the
abdominal cavity, this edge being really continuous with the
horizontal septum” (post-hepatic septum). This condition I
believe to be due to the fact that in the Emu the posterior parts
of the post-pulmonary septa fail to unite with the lateral body-
walls, as they do in most forms, but retain throughout life the
embryonic relations shown in text-fig. 29,p.217. Beddard proceeds
to say, “the oblique septum is thus merely a fold of the horizontal
septum; they form one continuous structure.” In the adult the
two septa are really continuous, but it must be borne in mind
that from its development only the anterior dorsal moiety of the
post-hepatic septum is of the same nature as the post-pulmonary
septum, the posterior ventral portion having a quite separate
origin.
The other modification of the relations of the oblique septa
occurs in many—and possibly all—Passerines. Here the septa
of each side, ‘instead of being attached independently to the
sternum, become fused with the falciform ligament in the middle
line, and form a horizontal sheet of membrane covering over the
two lobes of the liver. The original (?) attachments of the oblique
septa are not, however, in these birds entirely lost; a much
fenestrated membrane—sometimes, indeed, reduced to a thread
or two—remains to remind the anatomist of the more prevalent
conditions. In the Rook, however, they are completely preserved.
But the attachment of the falciform ligament to the sternum in
the middle line is lost.” This condition of the oblique septa
appears to be due to the backward growth, on the ventral side
between the sternum and the pericardium, of diverticula of the
median interclavicular air-sac. These in the adult seem to form
a single sac, but are presumably of paired origin, since they open
into the interclavicular air-sac by distinctly paired orifices; and
Bertelli [7] has shown that this air-sac itself arises in the embryo
as a double structure. These diverticula would seem to push their
way behind the peritoneum lining the ventral body-wall, and so
carry it inwards until it comes to invest closely the pericardium
and liver-lobes, as Beddard shows to be the case in the adult bird.
The persistence or not of the sternal attachments of the oblique
septa would in this case be of no especial significance, since it
226 MISS M, POOLE ON THE DEVELOPMENT OF THE _ [ Mar. 2,
would depend only on whether, as in the Rook, the diverticulum
remains comparatively slightly expanded laterally, and so only
detaches the peritoneum from the body-wall in the middle line;
or whether, as may be the case in other birds, it extends far
round against the lateral body-walls into the post-pulmonary
septa.
As I have before mentioned, I cannot agree with Bertelli’s
contention that the oblique septum, the « diaframma toraco-
adeomumells,” is not a septum, or part of a septum, at all, but
‘non é altro che la parete ventrale dei sacchi aeriferi intermedii e
la parete posteriore del pericardio.” He shows how in the Fowl
the ventral walls of the air-sacs are covered by muscular tissue,
and how in the Duck this is more markedly the case; but holds
that the muscular tissue has nothing te do with the subdivision of
the body-cavity, but belongs to the walls of the air-sacs, together
with the nerves which, according to Beddard, are of spinal
origin. Since, as I have show. the air-sacs never project
beyond the edge of the post-pulmonary septum, the muscles would
lie in the connective tissue between the peritoneum bounding the
abdominal cavity and the endoderm of the air-sacs, and therefore
in the tissue of the post-pulmonary septum. And I believe that
the presence of muscular tissue justifies the description of this
part of the abdominal wall as the postero-ventral portion of the
post-pulmonary septum.
Huxley [12] also describes a layer of muscular fibres in the
oblique septum of the Duck, and Beddard [2] finds a considerable
sheet of muscle in part of the septum of the Puffin (/ratercula
arctica). He says: “ This layer of muscular fibres arises from the
pubis—from the proximal and larger half of this bone; it is
abundantly furnished with blood-vessels and nerves. The direction
of the muscle is oblique ; it covers the hinder region of the oblique
septum, ending abruptly some little way in front of the posterior
attachment of the latter ; it is attached below to the upper surface
of the sternum, and to the abdominal parietes along the last
sternal rib.” Beddard also finds similar muscles in two species of
Penguins (Hudyptula minor and Spheniscus demersus) and also,
though feebly developed, in the Toucan (Rhamphastos ariel).
The Post-hepatic Septum.
This septum in the adult fowl isa membrane extending, as |
have shown, obliquely across the visceral cavity from its dorsal to
its ventral wall. Dorsally and laterally in front it is continuous
with the post-pulmonary septum, which in turn is united with
the vertebral column by means of the median dorsal mesentery.
Posteriorly and postero-laterally it is attached to the ventral
body-wall. It thus forms a complete septum, except on the left
side where the pulmo-hepatic recess remains in communication
with the abdominal cavity by means of a narrow aperture.
Except for this orifice, the two pulmo-hepatic recesses now form
1909. ] SUBDIVISIONS OF THE BODY-CAYITY IN BIRDS. 227
entirely closed cavities, lying between the post-pulmonary and
post-hepatic septa, the alimentary canal and the liver. The post-
hepatic septum passes dorsal to and behind the liver-lobes, and
ventral to and in front of the intestines, and carries on its ventral
surface the right anterior abdominal vein. The gizzard is enclosed
within its thickness, in the ventral portion of the septum, and is
thus completely shut off from the ventral liver-sacs, the posterior
walls of which are constituted by the anterior peritoneal layer of
the sheath of the gizzard.
This septum has been described in the Stork (Lepétoptilus) and
the Flamingo (Phanicopterus antiquorum) by Weldon {17 |, and in
Rhea, Struthio, and Dromeus by Beddard [1]. Inthe Struthiones
the liver-sacs are said to be closed off posteriorly from the
space ventral to the post-hepatic septum, and this may possibly
be compared with the condition found in the Crocodilia (see
below). Beddard also describes the septum in Cranes and Horn-
bills, in the Penguin (Spheniscus demersus), in the Spur-wingerl
Goose (Plectropterus gambensis), and in Bernicla brenta, where
the septum bears a second (left) anterior abdominal vein which
apparently corresponds to that occurring in reptiles, in the Grue
(Cariama) and in many other birds; and I have myself observed
it in the Golden Eagle. It probably therefore occurs throughout:
the entire group.
Ill. THe SrrRucTURES POSSIBLY REPRESENTING THE
AVIAN SEPTA IN REPTILES.
There are but few descriptions of the subdivision of the ecelom
in the Reptilia, and these deal mostly with the adult anatomy ;.
so that it is impossible to establish any homologies between these
forms and Birds, in the almost complete absence of embryological
evidence. I have therefore merely collected such facts regarding
the subdivision of the ccelom in the various groups as may be of
significance when more work on the development has been done.
In the Lacertilia generally there is no representative of either
the post-pulmonary or the post-hepatic septum, but in many
genera — Lacerta, Iguana, Chameleo, and others—there is a
well-developed pulmo-hepatic ligament, and consequently a pulmo-
hepatie recess is formed on the right side [9]. This arrangement
occurs again in Sphenodon, and is found of course also in
Mammals [13]. In the Teiidee, however, Butler describes a well-
developed post-hepatic septum in Z'upinambis teguexin, and a
slight approach to this condition in Ameiva surinamensis and
Callopistes maculatus, where the median ventral ligament is ex-
panded laterally behind the liver, but does not become attached
to the body-walls. In Zupinambis, however, the subdivision of
the body-cavity into a pulmo-hepatic and an intestinal portion is
almost complete except for a small aperture on either side. These
are situated in the dorsal moiety of the septum, and hence Butler
compares the latter with the ventral or omental part of the post-
hepatic septum of birds; the absence of the dorsal portion in
228 MISS M, POOLE ON THE DEVELOPMENT OF THE __[ Mar. 2,
consequence of the absence of ‘air-sacs being indicated by the
apertures in thisregion. That part of the septum which is present
dorsally, he accounts for by the need of some such transverse
membrane to carry the inferior vena cava to the liver, in the absence
of the pulmo-hepatic ligament present in most other Lizards; and
suggests that it may have been produced by the vena cava,
instead of developing as in other Lacertilia and Birds in tissue
continuous with the right pulmo-hepatic ligament, arising in a
lateral expansion of the median dorsal mesentery.
In many species of Monitor and Varanus, Beddard [3] describes
a horizontal (post-hepatic) membrane which covers the intestines
when the body-wall is opened ventrally. He says it is attached
anteriorly to the ventral body-wallin the middle line, and dorsally
to the spinal column, while to the lateral parietes it is attached
here and there by membranous bands. ‘It passes over the lobes
of the liver and the stomach and shuts off the lungs from the
abdominal cavity. The umbilical ligament dividing the two liver-
lobes is present, and is attached to the dorsal side of the hori-
zontal membrane. This horizontal membrane also separates the
kidneys from the reproductive glands; the latter lie internally to
it; the kidneys are placed outside it....The fat-body when
present lies below the membrane, and is therefore shut off from
the abdominal cavity.” Butler [11], however, describes how the
fat-bodies in reptiles are typically placed outside the ccelom and
surrounded by distinct extra-peritoneal cavities-—the circum-
adiposal spaces. He therefore considers it probable that the
greater part of this horizontal membrane described by Beddard
is the lining peritoneum which has become almost completely
separated from the abdominal walls by the great extension of the
circumadiposal cavities outside it. This view as to the nature of
Beddard’s horizontal septum seems to be further borne out by the
position of the umbilical ligament, which he describes truly as
being attached to the dorsal surface of the membrane; for in
birds and crocodiles, where I believe a real post-hepatic septum
exists, the ligament is attached to the ventral surface, and thence
passes to the ventral body-wall. Butler also shows that the
position of the kidneys and gonads in relation to the horizontal
membrane, is no proof of the septal nature of the latter; for the
kidneys are entirely excluded from the ccelom in crocodiles, birds,
chelonians, and sometimes in lizards, while they lie well within
the body-cavity in the Amphisbeenide; and in Monitor niloticus
the posterior part of these organs lies within, and the anterior
portion without, the peritoneal cavity. That part of the septum
which shuts off the lungs from the abdominal cavity is, however,
quite a separate structure, and appears to resemble more closely
the condition seen in some Chelonia than in any other reptiles.
The lungs of Varanus seem also to be so intimately connected
with the dorsal body-wall, that all trace of a pleural cavity is lost,
and thisagain happens in 7'’estudo (see below). Or Butler suggests
that the exclusion of the lungs from the peritoneal cavity may
1909. | SUBDIVISIONS OF THE BODY-CAVITY IN BIRDS. 229
take place in the Varanide as in snakes, by the gradual growth
of connective tissue from the ventral surface of the lung round
to the dorsal side so as to fill up any cavity that originally
surrounded it.
In the Chelonia, Bertelli describes the exclusion of the lungs
from the general body-cavity by the development of a septum
which he takes to be the homologue of the “diaframma ornitico.”
In the adult Testudo greca, this forms the ventral boundary of
the lungs-—round which it is fused with the lateral body-walls—
as a layer of fibrous connective tissue covered by peritoneum.
This condition 1s connected with that usually found among
reptiles by intermediate stages, seen in Vhalassochelys caretta,
where this layer is very thin and does not completely shut off the
lungs from the other viscera, and in Hmys lutaria, where it is
less developed, and the lungs for the greater part project free
into the pleuro-peritoneal cavity. In an embryo of Testudo,
32 days after oviposition, the relations of the lung rudiments and
the dorsal and ventral pulmonary ligaments are essentially the same
as those in a chick of 72 hours’ incubation (text-fig. 24,p.212); and
for some time development proceeds in the same way as in a bird,
the lungs extending laterally and dorsally, the growth in the
former direction tending to close off the pleural from the rest of
the peritoneal cavity, and that in the latter to reduce the size of
the pleural part of the celom. Later, the thick layer of tissue on
the ventral surface of the lungs, which separates these organs
from the underlying liver, meets and fuses with the lateral body-
walls, and thus constitutes an almost complete septum across the
peritoneal cavity. At the same time, the lungs unite with the
dorsal and lateral walls of the pleural cavity so as to obliterate
the ccelom in this region.
The diaphragm in 7'’estudo is thus formed, according to Bertelli,
out of the same constituents as is the diaphragm of birds, and has
the same position and relations. Also the gradual obliteration of
the pleural cavities takes place in the same way in the tortoise
embryo as in the chick ; although in the latter it is never com-
plete, and there remains throughout life a narrow but distinct
cavity bridged across here and there by strands of connective
tissue. Since, however, the growth of the lungs in a dorsal and
a lateral direction goes on at the same time, the obliteration of
the cavity round the lungs, and the completion of the diaphragm,
take place practically simultaneously; so that by the time the
diaphragm is constituted in order to close off the pleural part
from the rest of the peritoneal cavity, no pleural cavity remains.
It appears, therefore, to be more likely that the condition seen in
Testudo is to be compared with that occurring in the Varanide
rather than with that found in Birds.
From the structure of several Crocodiles which I had the
opportunity of examining, I believe that the arrangement of the
ccelomic septa in this group of reptiles approaches most nearly to
the avian condition, as Huxley and Beddard have already pointed
'230 MISS M. POOLE ON THE DEVELOPMENT OF THE _ [{ Mar. 2,
out [1, 2,12]. Here the post-hepatic septum is well developed, and
passes from its anterior dorsal attachment to the vertebral column,
behind the liver-lobes and pericardium, to unite posteriorly with the
ventral body-wall. As in birds, it encloses the stomach within its
thickness, and carries on its ventral surface the anterior abdominal
veins to the liver. It is attached to the lateral body-walls
throughout its extent, so as to separate completely the pulmo-
hepatic from the abdominal portion of the celom. Butler describes
in a very young specimen a small passage between the abdominal
cavity and the right pulmo-hepatic recess; but this presumably
becomes closed at a later stage, for there was no such communi-
cation in the other animals which I dissected. The post-hepatic
septum is apparently split secondarily in its ventral portion, and
the flap thus formed on each side is closely applied to the liver-
lobe so as to form a posterior wall to the liver-sac, and—together
Text-fig. 35.
LP. hp. spt.
\
eNO
Diva GT /
ERP St. Php.c. Ben
Young Crocodile bisected in the same way as Chick in text-fig. 34, p. 223.
with the oblique ligament of the liver with which it is connected
—to shut off the pulmo-hepatic cavity from a subseptal, ventral,
post-hepatic space. There is a large pulmo-hepatic cavity extend-
ing right round from the dorsal attachment of the post-hepatic
septum to the ventral attachments of the oblique ligaments of
the liver, perfectly continuous except for a narrow partition
projecting a little way inwards from the ventral body-wall. ‘The
pulmo-hepatic recesses seem to correspond almost exactly to those
spaces in birds, lying as they do between the ventral surfaces of
the lungs, the dorsal margins of the liver, and the post-hepatic
septum; and I believe that tissue which forms the roof of each
recess, and covers the postero-ventral face of each lung, represents
the post-pulmonary septum of birds. Text-fig. 36 is a dissection
from the ventral side of a young specimen of Crocodilus palustris.
It shows the continuous pulmo-hepatie cavities closed off behind
1909. | SUBDIVISIONS OF THE BODY-CAVITY IN BIRDS. 231
by the oblique ligaments of the liver and the more vertical portion
of the post-hepatic septum. Nearer the middle line lie the
ventral liver-sacs between the posterior wall of the pericardium,
the oblique ligaments of the liver, the post-hepatic septum, and
the median ventral mesentery. Behind, the post-hepatie septum
Text-fig. 36.
Young Crocodile opened from the ventral side. The ventral walls of the peri-
cardium, pulmo-hepatic cavities, and ventral liver-sacs have been cut away.
lies horizontally over the stomach, attached both posteriorly and
laterally to the body-wall (¢f text-fig. 32, p. 222, ofa bird), Text-
fig. 35 is of the same Crocodile bisected longitudinally a little to the
left of the middleline. It shows, in addition to the structures seen
232 MISS M. POOLE ON THE DEVELOPMENT OF THE [ Mar. 2,
in the previous figure, ‘the incomplete ventral partition between
the pleural and hepatic regions of the pulmo-hepatic cavity, the
position of the left pulmo-hepatic recess, and the possible repre-
sentative of the avian post-pulmonary septum (cf. text-fig. 34 of a
bird bisected in the same way, and also text-figs. 33, p.223,and 37).
Text-fig. 37.
Phrs. Ppu.spt.(?)
as BGS ene SO Rose ete —— ote 4
Php.c. Kilizs. NN ‘Po, Llu.lpo
Ob.lg.lv.
Diagrammatic plan showing the subdivisions of the ccelom in a Crocodile
in longitudinal section.
Huxley first called attention to the similarity between the
arrangement of the ccelomic subdivisions in Crocodiles and in
Birds. He describes a fibrous expansion extending from the
vertebral column ‘over the anterior face of the stomach, the
liver, and the dorsal and front aspect of the pericardium, to the
sternum and parietes of the thorax, separating the thoraco-
abdominal space into a respiratory and a cardio-abdominal cavity,
and representing the oblique septum of the bird. ... A broad,
thin muscle arises, on each side, from the anterior margin of the
pubis; and its fibres pass forwards, diverging as they go, to be
inserted into the ventral face of the posterior part of the peri-
cardium and into the ventral and lateral parts of the fibrous
capsule of the stomach, passing between that organ and the
adherent posterior face of the liver, and being inserted into the
fibrous aponeurosis which covers the anterior face of the stomach,
and represents the oblique septum.” This description appears to
refer to the whole of that membrane which I have called the
post-hepatic septum, together with the roof of the pulmo-hepatic
recesses and the oblique ligaments of the liver; and as I have
already said, it is only that tissue forming the roof of the pulmo-
hepatic recess on each side which I regard as possibly comparable
to the oblique septum (post-pulmonary septum) of Birds.
On the other hand, Beddard [2] says with reference to the
muscles described by him in the oblique septa of Puffins and
1909. | SUBDIVISIONS OF THE BODY-CAVITY IN BIRDS. 233
Penguins, that the muscle referred to by Huxley in the Crocodile
is the equivalent of this muscle, for in both cases the fibres arise
from the pubis. Beddard therefore considers it more justifiable
to identify the middle portion of the horizontal septum of the
Crocodile with that structure in birds, while the lateral portions
containing the muscles he compares with the oblique septa of
birds. And this comparison he considers is made more obvious
by the relations of the two septa in Dromeus; for here the oblique
septa are not attached to the ventral body-wall, but posteriorly
project with a free edge into the visceral cavity, appearing as a
fold of the horizontal septum. This peculiarity in the Emu
Beddard takes to support his contention that “the omentum
(horizontal septum) as well as the oblique septa of birds are to be
derived from the fibrous expansion which covers over the viscera
in the Crocodilia.” He also suggests that possibly “the oblique
septum of birds has been produced by a vertical fold of this ex-
pansion.” This does not seem to me likely however, as in
development it is the oblique septa which are first formed, and
only much later does the horizontal septum arise, partly as folds
of the former. Also, against the above comparison of the lateral
parts of the crocodilian horizontal septum with the oblique septa
of birds, should I think be set the fact, that the only cause of
the posterior attachment of the oblique septa in the latter being
so far back as to bear a resemblance to the position of the lateral
parts of the horizontal septum in Crocodiles, is the growth of the
posterior air-sacs. But since these structures are not present in
reptiles, we should expect to find the post-pulmonary septum but
little extended, and comparable more or less to the embryonic
condition in a bird (¢f. text-fig. 25, p. 213) before the air-sacs are
developed. And this is what we do find, if we take only that tissue
on the ventral face of the lungs in the Crocodile as representing
the avian post-pulmonary septum.
As to the origin and phylogenetic significance of the subdivisions
of the pleuro-peritoneal cavity, it is impossible at present to say
anything definite; for while on the whole the condition in the
Crocodilia seems to approach most nearly that in Birds, yet the
manner of the exclusion of the lungs from the peritoneal cavity
in the latter is far more like that which occurs in TVestudo and
probably also in the Varanide. While again the post-hepatic
septum—not represented in these two forms—is found, in the
same way as in the Crocodile, in the Telide, where it appears
perfectly comparable to the more ventral portion of the avian
post-hepatic septum; and according to Butler, it is also com-
pletely represented in Snakes. Therefore, whether or not the
subdivision of the celom will prove of importance as to the
phylogeny of the Reptilia and Aves, must, I think, remain
undecided until further embryological work has been done.
Proc. Zoou. Soc.-——1909, No. XVI. iG
234 THE SUBDIVISIONS OF THE BODY-CAVITY IN BIRDS. [{ Mar. 2,
List of Works referred to in the text.
1. Bepparp, F. E.— Notes on the Visceral Anatomy of Birds.
—I. On the so-called Omentum.” Proceedings of the
Zoological Society, 1885.
3. Bepparp, F. E.—‘ Notes on the Visceral Anatomy of Birds.
II. On the Respiratory Organs in certain Diving Birds.”
Proceedings of the Zoological Society, 1888.
3. Bepparpb, F. E.— On certain points in the Visceral Anatomy
particularly of Monitor.” Proceedings
of the Zoelosiea Society , 1888.
4, Bepparp, F. B.—* On the Oblique Septa in the Passerines
and in some other Birds.” Proceedings of the Zoological
Society, 1896.
5. Bepparb, F. E.—The Structure and Classification of Birds.
London, 1898.
6. Berretir, D.—‘‘ Contributo alla morfologia ed alla sviluppo
del diaframma ornitico.” Monitore Zoologico Italiano,
ix. Firenze, 1898.
7. Berreitt, D.—‘‘ Sviluppo dei sacchi aeriferi del pollo. Divi-
sione della cavitd celomatica.” Memorie della Societa
toscana di Scienze Naturali, xvil. Pisa, 1899.
8. Berretti, D.—“ Ricerche di Embriologia e di Anatomia
comparata sul diaframma e sull’ appar ecchio respiratorio dei
vertebrati.” Archivio di Anatomia e di Embriologia, iv.
Firenze, 1905.
9. Buruer, G. W.—*‘ On the Subdivision of the Body-cavity in
Lizards, Crocodiles, and Birds.” Proceedings of the Zoo-
logical Society, 1889.
10. Butter, G. W.—‘ On the Relations of the Fat-bodies of the
Sauropsida.” Proceedings of the Zoological Society, 1889.
11. Butter, G. W.—‘‘ On the Subdivision of “the Body- cavity in
Snakes.” Proceedings of the Zoological Society, 1892
12. Huxtey, T. H.—‘On the Respiratory Organs of dApteryx.”
Proceedings of the Zoological Society, 1882.
13. Maui, F.—‘“ Development “of the lesser Peritoneal Cav ity in
Birds and Mammals.’’ Journal of Morphology, v. 1891.
14. Miituer, B.—‘‘ The Air-sacs of the Pigeon.” Smithsonian
Miscellaneous Collections, Quarterly Issue, iv. 1908.
15. Rocust, G.— Contribution 4 Pétude de Yanatomie comparée
des réservoirs aériens d’origine pulmonaire chez les oiseaux.”
Annales des Sciences naturelles, ser. 7, x1. Paris, 1891.
16. Saprpny, P.—Recherches sur Jlappareil respiratoire des
Oiseaux. Paris, 1847.
17. Weupon, W. F. R.—‘“On some points in the Anatomy of
Phenicopterus and its Allies.” Proceedings of the Zoolo-
gical Society, 1883.
See also Hoehstetter, F.—‘* Ueber die Entwickelung der Scheidewandbildung in
der Leibeshéhle der Krokodile,” in Voeltzkow’s “ Reise in Ostafrika,” 1903-1905,
Bd. 4; and “ Beitriige zur Entwicklungsgeschichte der Himys lutaria. 2. Die ersten
Entw icklungsstadien der Pane und die Bildung der sogenannten Nebengekrase,”
in the Denkschriften der Math.-Naturwiss. Klasse der Kais. Akad. der Wiss., Bd. 84,
Wien. 1908, which I have been unable to consult.
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SHETES) On PATE TEA VWIlEGATAS <7
Bale & Damelsson .L'¢
1909.] THE SHELL-GROWTH OF THE LIMPED PATELLA VULGATA. 235
EXPLANATION OF ABBREVIATIONS IN TEXT-FIGURES.
Abd.c. Abdominal cavity. | P.hp.spt. Post-hepatic septum.
A.int.as. Anterior intermediate air-sac. | Ph.rs. Pulmo-hepatic recess.
Cf. Line of fusion of celomic epi- | P.int.as. Posterior intermediate air-sac.
thelium of post-pulmonary | Pl.c. Pleural cavity.
Septum with that of lateral | PJ.hp.c. Pulmo-hepatic cavity.
body-wall. | Plt. Pulmonary rudiment.
Ct.pl.m. Costo-pulmonary muscles. P.pl.spt. Post-pulmonary septum.
D.ms. Median dorsal mesentery. P.p.p.m. Pericardio-pleuro-peritoneal
D.pl.lg. Dorsal pulmonary ligament. membrane.
z. Gizzard, Pin. fd. Pronephric fold.
Ht. Heart. | Pt.c. Peritoneal cavity.
Kd. Kidney. | Spl. Spleen.
Ing. Lung. | WSp.rnl. Supra-renal body.
Lv. Liver. | S¢. Stomach.
Mi.dé. Miillerian duet. Um. Umbilical opening.
Ob.1g.lv. Oblique ligament of the liver. | V.lv.s. Ventral liver-sac.
(Hs, (Esophagus. Pms. Median ventral mesentery.
P.as. Posterior air-sac. | V.pl.lg. Ventral pulmonary ligament.
Pe. Pericardium. | Wy.by. Wolttian body.
P.hp.c. Post-hepatic cavity.
2. The Growth of the Shell of Patella vulgata L.
Biv 1d, SS). Levorsetaore, Wilk
[Received February 16, 1909. ]
(Plate XXXII. +)
This paper consists of two parts; the firsta study of the rate of
growth of the lmpet-shell, the second a study of the gradual
change in the shape of the shell as it grows in size. Throughout the
paper the size of the shell has been taken as a measure of its growth.
Owing to the manner in which growth takes place, no dimension
of the shell can increase without a corresponding increase in
the other dimensions, so that any one dimension may be taken
as a measure of growth, 7. e. of increase in size. It is convenient
to take length as a measure of size. Knowing the length one can
calculate with a fair degree of accuracy all the other dimensions,
if one has previously determined the average ratios of the other
dimensions to length for successive values of the length.
I. Rare or Grows.
1. Breeding Season.
The fact that limpets of 5 mm. in length may be found in
January and in July leads one to suppose that the breeding
season is of considerable duration, and the observations which I
have made fully confirm this opinion.
I tried to discover at what time of year small limpets chiefly
appear, by watching events in the small bay where I studied the
growth of the limpet. On January 11th, 1908, there was a
** Communicated by Prof. J. ARTHUR THomson, F.Z.S.
+ For explanation of the Plate see p. 253.
16*
236 MR. E. 8. RUSSELL ON THE SHELL-GROWTH [Mar. 2,
small proportion of 10-15 mm. shells and on January 25th there
was seemingly a greater proportion of small shells 7-10 mm. in
length. After that I have no records which indicate the settling
down of very young limpets until July 21st, when I found that
the small shells were mostly 15-25 mm. in length (and therefore
simply the descendants of the 10-15 mm. long limpets of
January), but that an occasional small one (8 mm.) occurred
among them.
On August 4th at Millport I collected one or two tiny shells
2°5-3 mm. long from the crevices of a rock in the lower half of
the beach. On September 2nd at the Cloch, Gourock, there were
visible in the bay a few 5-10 mm. shells, and on October Ist there
were a few shells 8-12 mm. in length in places where they were
not present on September 2nd.
From these data it would appear that the breeding-season ex-
tends from July to December or January. I examined also the
gonad of a number of limpets at various times during the year.
On September 18th, 1907, three limpets were examined. ‘Two
of them were females, 28 mm. and 40 mm. in length, and the
ovaries contained large but not quite ripe ova. In the 40 mm.
long male the sper matozoa were fairly ripe. A number collected
at the Cloch on December 25th, 1906, were quite ripe. Of 26
limpets collected at Cardwell Bay, Gouroek, on December 28th
and 31st, 1907, 21 were males and only 5 were females. The
males were not very ripe and some of them seemed spent,
but the females had mature sexual products. Of five males
collected at the Cloch on January 11th, 1908, only one was at all
ripe, the others being spent. Ten limpets, 4 females and 6 males,
were taken on February 8th. The males seemed spent and the
ova in the gonads of the females were disintegrating. The
breeding season is clearly over by the end of January if not a
little sooner.
The sex of six limpets of 35-42 mm. in length collected on
the 23rd May could not be determined with certainty, though
they appeared to be males. The gonad was brick-red and only
one-half to three-quarters its full size. At the anterior end it
was beginning to turn cream-coloured, as if it were ripening into
a-mature testis in that region.
A male examined at Kames on July 2nd, 1906, had well-
developed spermatozoa.
On July 11th, eleven limpets were examined. Six of them
were females and their ovaries were at all stages of ripeness. One
or two showed a number of fairly ripe ova, and the ovar y had the
typical olive-green tint which it shows when at all mature. The
others were “extremely unripe with very minute ova, and the
colour of the ovary was a reddish-brown. The males were all
moderately ripe. J attempted to carry out artificial fertilisation,
and rather to my surprise obtained a number of early segmen-
tation stages.
Artificial fertilisation was carried out with success at Mullport
on August 4th, early segmentation stages being again obtained.
1909. | OF THE LIMPET PATELLA VULGATA. 237
Of 23 limpets collected at the Cloch on October Ist, 19 were
males and fairly ripe, 3 were females also ripe, and 1 was imma-
ture, being only 20 mm. in length.
The breeding-season, therefore, probably lasts from about the
middle of July to the middle of January. There is much in-
dividual variation as to the time of ripening, and so it is
improbable that any single limpet is mature throughout the
whole breeding-season. While the commencement of the
breeding-season seems to be ill-defined, its finish is mere
definitely fixed.
Dr. J. F. Gemmill, who has examined the gonads of many
limpets, tells me that he considers the limpet to be ripe from the
beginning of November to the first fortnight of January. Davis
and Fleure write (2. p. 59), “the season of sexual maturity is
the autumn ; Boutan finds it to be about September at Roscoff ;
at Aberystwyth we think it is somewhat later.”
According to Fischer (3. p. 867) “la ponte s’effectue sur les
cotes océaniques de France a la fin de mars et au commencement
davril,” which agrees well with the observations detailed above.
It is probable that on our own shores the limpet is ripest during
the months of October, November and December. <A. similar
state of affairs seems to hold in the Adriatic, for Patten (8)
obtained artificial fecundation of the eggs of Patella at Trieste
in the beginning of December and the middle of January. At
Naples, however, Wilson found the eggs of Patella cwrulea in a
mature state from March until June (12. p. 199). |
Size at which maturity is reached.—From the examination of a
large number of limpets (80—-90)'T find that sexual maturity is
reached by the males at a length not less than 21 mm. I have
found immature males during the breeding-season 16°5, 19°5, 20,
and 21 mm. long. The gonad grows with the animal, and in
males 21-30 mm. in length it is only about half its ultimate size.
It is fully developed at about 40 mm.
I have seen no female smaller than 26 mm., but at that length
the limpet may have mature ova. The ovary attains its full size
in limpets of about 40 mm. in length.
According to Dr. Gemmill’s observations limpets may be
mature as small as 18-20 mm.
It is of interest to compare the closely allied Acmea with
Patella in this respect. In Aemca (the breeding-season of which
lasts from the middle of April to the end of July) sexual maturity
is attained under 10 mm. long and probably after one winter
(Willcox, 11).
Another interesting example from among the Gastropoda of a
very rapid attainment of sexual maturity long before adult size
is reached is to be found in Semper’s ‘ Animal Life’ (p. 126).
Land snails in the warm regions round the Mediterranean are
brought to sexual maturity when only six months old by the heat
and moisture of the spring. They are then by no means full-
grown.
Proportion of the Sewes.—Out of 87 limpets examined, 66 were
238 MR. E. 8. RUSSZLL ON THE SHELL-GROWTH [Mar. 2,
males and 21 females. They were collected for the most part
near Gourock. The males are apparently about three times as
numerous as the females in this locality, while at Muillport
Dr. Gemmill (5. p. 394) found that of about 250 limpets examined
68 per cent were females. From some old observations by
Lebert and Robin (6) it appears that in their experience there
were about three males to eight or ten females, It would be
interesting to discover the reason for this great variation in the
proportion of the sexes.
2. Growth Data.
In order to study the rate of growth of the limpet throughout
the year I marked a considerable number on December 31st,
-1907, in a bay near the Cloch, Gourock, and attempted to keep
full records of their growth from month to month during 1908.
A number of them of course disappeared during the year, but
their places were filled by others; for example on May 25th and
May 30th, I measured about two dozen small-sized limpets and
kept many of them under observation until the end of the year.
The detailed records are given in Table I. where they are
arranged in year-groups, and more or less in order of magnitude
in each group. M = mean value.
The possibility of arranging the measurements in year-groups,
at least for the first two year's, arises from the fact that the
breeding-season of the limpet, though a very long one, is fairly
well-defined.
Taste I.—Growth-Data, arranged in Year-Groups.
1908 brood,Oct. 31, 11-5 mm. Nov. 30, 13 mm.
Frrsr Year-Grovur (1907 brood).
1907|1908 | | | |
\Dec.|Jan.|Feb.|Mar. Apr.) May May|May June] July |Sep. Oct. | | Oct. Nov. Dee.
SUS WEG, | Lan (ose) 5s a0) 27. |) 80: ari) ee soe
| eee ee | F uw | —|—
Tad oe (fox (OS VOR ees |e |G eka
Pes) cea. 0 || \aa9! a4e5 | | |
enoleEy 10-4) 11 | 19:5) 15°3) | .. | 189 | 19°8 | 23°7 26 27-6
1 PG Gos ines ina 168 = ee ee 238 | 271 Bl elena
5... 14) 12 | ... 164... | .. 20892626398 [o03) |
Bhi | ae ies 9 Fire ae |
his | fea ea) tay 20291 21
eae ho. | lig |... 15 |171/20 |22 |23 apm
9... be ne 119 149 | 182 | 208 225 Sdawillinies
Teta Ba ee ard iowileemn | Nees lila
1909. | OF THE LIMPET PATELLA VULGATA. 239)
First Year-Grour (1907 brood) (continued).
1907 1908 | lee
Dec. Jan. Feb. Mar. Apr.|May May|May| June | July | Sep. Oct. Oct.|Nov. Dec.
2h 745)5|| tek |) eh, | IG) I 25) GOL. ge || XO, | 2 |, || Gal) BIO.) Sil,
| | |
oe = | | | |
Hilal ee | “ | sop, |) soe Ih ome l apace PMO) sce Tee, 20." | | |
y |
West see ll soe | oe) een ll coe Wh aus) ces 1D 1) aiGe | sHaRe, | ones! Daca
||}, —
1B} ee 2 (aa" | aes es) aire oars\benleralars
Wc ose) coont abe dl accel) see || eesa tM Meme IG RN SUSHS | aapd) oF
) 15.) |. |. |. |. | [44]... | 16-4 | 191 | 21 | 23-4] 24:5] 96-9] 27
[16 .| |e |e oe | |. | |14t8] 166 |17 | 20. | 289) 24-5) 245) 247)
14 | 169] 21 |24 | 26-4 97-9/ 97-8] 28
|
8...) 2 Jaga) | 69 | 20°9\| aara| 25:3) 274
1d ell cal ool eal CE ee ee eran (245 27°8 29 | 29°8| 30
an | ale | 2 eee | S94 21-3 art oe
oe ee ee er ee ee es) Etza 21 245 26-4 28
Issel onl HG Stl 161 185 22 (25-4! 26°5 27-9, 28°7| 28-9
23) eG eae eee 16 [19 |21°5 | 24-7| 268129 | 2901299
Pa ee a ee aa 18 19°2 | 22°8 | 25 97-5 28-9 29-11 29-7)
25 | eee Geel des, de a 164 .. |195 [23 | 266 278 285 29°2| 29°
o8../ lel | Seles lasclas |
| 27 eS ail cooler 197 | 24 | 26-1 285 29 9,305 31
28 eal eee Tal. |ao9) 21-1 |99:1/93 | 932/95 | 25
Pie il len |... 196... |22 | 25-4 [a7 | 28 1) 28:8] 292] 29-8
20.) | | | | ES acer ailes ae (eoe) |
si al Cae | 1198 235 27-2 305/328 33-1 B38 338,
| 3) le ea ava |15:2 113 [Se ee
| 83... -, |142) awe 167 189 .. |... | 217/252 /aeelat |sea aes]
eo ol alee en al es ST a Gap |
Baal acl ao ha ae ee ee jesilgeilora| |
36 Os oa Hae | Nigeai ee eee 995/309, |
| a7...|... | ... |a7e| 182! ise\22 | eal He cone gee
| 38. Al 188 199 21 |. | .. | 245/278 |208) 316... |... |aaral
| 39.. Pa ..| ae fat [225] | 253 | 287 29°5| 3061 |
j W.-H
SEI Saher see eal | 15°5 186 | 21-2 | 24-5) 26°7) 27°8) 28:4, 29°1.
240 MR. E. S. RUSSELL ON THE SHELL-GROWTH [ Mar. 2,
SECOND YEAR-GrRovr (1906 brood).
| | i
| | 1907 | 1908 | : HASPR nee
| Dec. | Jan. |Feb.|Mar./Apr. May’ June | July |Sep.| Oct. Oct. Nov. Dec.
| | 31. | 1. Ee / Me 1d 28. 27. | 30. | 2.) 1. | 81.) 30.) 31.
“40 betel love 22°2| 23:1'23°6 267 | 286 | 30°4|30°8 324 32°5 33°5
41... | 218/221 22293 243 262) 27 28) ||
42... | 937/251 255 262.266 278) 293/30 |31
43 ...| 22 | pero ea | Bat cm Me ee er
| 44 | 295 ea 23 24-1 25°6 268 291 31-2 saat | 353)
45 ..| 238] .. loeajoue | | | tri al
nse) | eneies (269/272 284 298 | 329 35636737 37 37
47 || 27 |... [27 jars 282 29 | 308 33. | 3dr2/ 34°8 352 352 353
| 48...) 27-9 Cains .. (eee) | Diao sae Adee Wiese
49 .. | 27-2 ‘ar7| 23 28°5 28'5 30-2 | 308 31 | 312 BB 33°38 336
50.../ 29 | ... [99 /30 [a1 lara) sie| 34 [36 [36 [a7 |37aae
S| ees 291]... | .. {82 | Be | on 379 39°5 40 pee
a aoe | Nee! ay | ae a a a
53 s| set 299 50 es Patil eae Gee |
54 ...|| 30 poe [hes | ... |829) 84 | 369 38389380
55... 30 ee OR es eeu sieea "a7 B92 405 ae | 42-2) 40:2)
56 | 308 .. [a1 |... [82 |a20l ae9 | 35 ls7-8139 |30 [39 39 |
57 | ee 7 32'3| 32°8| 33-6 845, 37-4 | 40-8 (42 42 (42 42 | 422)
68... .. | aaa (328 a36|84 |35 | 37-6 01 409] | al
69. nannies 33 |s3 [333/34 ape bens Wile raat |
60) ...|| 32°6 | ee es ape ela bo) |
Merde ome Cer de oP wi aoe Sime ie
ellee loa ay ala Cenc wale ak ab |
63 | sea] 2 [359] faa || 4, irate
64 Pl) eeel Gee] 2 [aos 41 [41 [4 [41
65 Ll aN ad ie ae eae Se 40°6 408 41 41 |
- | 7 | | oo
M .../| 28:3] ... | 28:6| 984 29°6 311) 32:7 | 33°7 (36 |37-1) 38 |38 | 38
* No 51 measured 23°5 mm. on 7th Sept. 1907.
1909. | OF THE LIMPET PATELLA VULGATA. 24]
THirD YEAR-Grovur (1905 brood).
1907/1908) | | leg rot
| Dec. Jan. Feb. Mar. Apr. May June July Sep. Oct. Oct. Nov. Dec. |
| BL | 2 | 8: |iae | 1a.) 28. | 27. | 30. | 2. | 1. / 81. | so. | 31.
SE) (ase acelge Snel eae | al
66 i) B67. SSP eevee oe | 438 4d 45 |
67 al Seay enemlara0 Pie aaa ihe
68... 367 .. 369)... [372879 389 389 38998959 40 |40.
69...) 380| .. |. |39 |a96|40 | 41 | |
l7o..| . | a7 |.. larslars| ra dao a
ies a SINE Sil) Cl Ae le ae aa ae aaa
ell a [ocala ce ee Pass |
ral) ee 7) all ea a Ra 4a 43°5 43°9 43-9] 43-9
| | |
FourtH YEAR-Group (1904 brood).
|1907| j1g08! | e donne
Sept. Dec.| Jan. Feb. Mar. Apr. May! June) July | Sep.| Oct.| Oct. Nov.) Dee.
fle 1) Sil | lik) Gy | ily PEER Al MA Os | 25 ile |) Sik | BOs NS,
=a A a RR hos
74...) 435 468 46'8 47-1 47-4 | a
75.) = |43 |. [aol aa [aa 1) pe a es aay
76 “43°6 46 46 |46 | 46 | 47 | 6479 48 48 481 481
77 ...| 457 |46 |. |48 | 46 | Che tae oan ae
| FB cos | i545) | 455) | |
iva .../465/..| .. |a7 lar |are larva al
| 80. & Heh a 47-4 ATS ATS 47-5) 47-9)
| 81 Baal: ae 49°2 492) 493 493
|
Firrg or Sixra Year-Grovup (1903 or 1902 brood),
| 82 | 50 | 50 |50 | 50 | |
50 S05. |50°5| 51 514] 52) 52-2 | 52'5| 525 58 535 535
c |
first Year-Group.—It is a curious fact that one rarely finds
on the shore limpets less than say 7 mm.long. I have found them of
242 MR. E, 8. RUSSELL ON THE SHELL-GROWTH [ Mar. 2,
5 mm., 41mm., and 2°5 mm. in length, but such small limpets are
distinctly rare. They are of course extremely inconspicuous. In
the autumn and during the winter limpets of about 10 mm. long
gradually appear on the shore, settling down in greatest abun-
dance in the months of January and February. There seems no
reason to doubt that such limpets are only a few months old, and
represent the first year-group. As the tables show they grow
very rapidly (see Tables I. and V.) and can have taken only a few
months to reach the length of 10 mm. It should be noticed that
in very small limpets (2
in proportion to the animal than in older limpets. If these
limpets of about 10 mm. length represented not the first but the
second year-group there ought to be found on the shore in spring
and summer great numbers of shells from about 3 mm. to 7 mm.
in length, and this is quite certainly not the case. Nor does it
seem likely that the free-swimming larval life of the limpet is
very long, or indeed longer than a week or two, for in the allied
Acmea virginea the pelagic life is limited to a few days (Boutan).
Acmea testudinalis, a much smaller shell than Patella vulgata,
breeds from April to July and the season’s young are 4-5 mm.
long in October (Willcox, 11).
A random sample of 1003 limpets taken in July 1903 and
1904 at Southend, Arran, show very clearly that this group of
shells about 10 mm. long in January is really the first year-group.
The sample was taken just at the very beginning of the breeding-
season so that only a very few of the season’s young were
included.
Tasxe IT.
Random Sample in month of July.
Size in mm. No.. | Size in mm. No.
| 3— 55... S| ESOS SiO met cee ao |
6— 85... 9 | 39—41°5 50
| 9—115 ... 26 | ADI sas spose 65
VO VAS ii eeceedes 72 | 45—47°5 91
| lit W/o 9 ae 102 | 48—50'd 83
US =—PXORS caossecon ose 97 || ISS BPS) son ccsnscsns 77
PPE TSY saanneeoeone 55 | 54—56'5 ......-..... 39
DIM OIS scocaacstees eeu Whe “SISOB ssersccnan TD |
[eye ee Gt ORE he se |
[BOSS FS ccrssectache FAT Mas lh BSG an ta 1
ly UBB ea SB io uc wile tsa! eumnCG = 6b; 5 seme nites amma
The first year-group revealed by these figures has its mode
clearly in the fifth class (15-17°5 mm.). Leaving out the first
class (8-5°5 mm. shells), which represents probably the first of the
season’s brood, and taking the next seven we find that tne mean
value is 17°53 mm., indicating that limpets of the last season are
1909. ] OF THE LIMPET PATELLA VULGATA. 243
now in July of an average length of 17-18 mm. These facts
taken together seem to Vena us in placing in the first year-
group the limpets, Nos. 1-5 (Table I.). These reach a length of
about 15 mm. by the third week in May. Nos. 6-31 form a
natural group, since they were all found in a small area of a few
square yards and present no abrupt difference in Sse. Nos. 6-23
are quite clearly of the same year-group as Nos. 1—5,and it seems
safe to include also in this first year-group Nos. 24— 31 and 32-36.
No. 29 for example, though 19-6 mm. long on May 25th, is only
29°2 mm. long on Dec. 31st, a length which is reached on that
date by No. 25 which was only 16-4 mm. long on May. 25th.
One must allow for a good deal of variation in the rate of growth
as well as for the fact that each year- group may include limpets
differing as much as six or seven months in age, since the breeding-
season is of that length. Nos. 27 and 28 illustrate how one
limpet may differ from another in rate of growth, so also do Nos. 3
and 4. On the other hand the rate of growth often exhibits
surprisingly little variation, as is shown by Nos. 12, 13, and
14, 17 and 18, 24 and 25, and others.
The placing of Nos. 37, 38, and 39 in the first year-group is a
little doubtful, but is I think on the whole justifiable.
The mean values have been calculated from the month of May
onward, and from them we can infer that a limpet which has
come into existence say In September will reach a length of about
15 mm. in the following May, 18 mm. in June, 20 mm. in July,
24 mm.in August, 26 mm. in September, 27 mm. in October,
28 mm.in November, and by the end of the year will attain on an
average a length of 29 mm.
These figures are of course only roughly approximate and are
valid only for the particular locality where the limpets grew.
Limpets at the Southend of Arran seem to reach a length of only
17-18 mm. by the month of July.
Second Year-Growp.—Given the first year-group it is a simple
matter to determine the second year-group. On December 31st,
1908, the first year-group had a certain range and a certain mean
value, and the second year-group must necessarily have had on
December 31st, 1907, approximately the same range and the
same mean value. The limpets assigned to the second year-group
(Nos. 40-65) have been selected on this principle. Their range
of size on December 31st, 1907, 1s from 22 mm. to 34:4 mm.,
and their mean 28°83 mm. The mean value of the first year-
group on December 31st, 1908, was 29°1 mm. and the range from
24:5 mm. to 33°8 mm., but if No. 7 had survived till December
31st the range would have been from about 21:5 mm. up to
33°38 mm. Theaverage size on December 31st, 1908, in this year-
group is 38 mm.
Subsequent Year-Groups.—lt is not possible to arrange the
limpets of 35 mm. long and upwards in year-groups with any
certainty. The arrangement adopted above seems the most
satisfactory, but it is only tentative.
244 MR. E. 8. RUSSELL ON THE SHELL-GROWTH [ Mar. 2,
3. Variation in Rate of Growth with Increase in Size.
The rate of growth decreases with increase of size. This fact
is shown by the data already given, but in order to gain a precise
knowledge of the manner of the decrease it is necessary to dis-
count the possible influence of time of year and temperature upon
the rate of growth. This has been done by taking the figures
given in Table I., for the period July 30th to September 2nd, and
considering along with them some other additional data which
have reference also to the month of August. These are,
(1) observations on the growth of 10 limpets at the Cloch,
Gouvock, from the end of July to the second week in September
1906 (43 days), and (2) on a similar number at the same place
from July 31st to September 7th, 1907 (38 days). <A few
additional data referring to seven limpets at Kames, during July
1906, are also incorporated. The detailed observations are given
in the following Tables.
TasxeE III.
Growth during 43 days. The Cloch, Gourock, Aug. 1906.
| |
is | weft By 2 : Growth per cent.
Group. Original size. = Final Size. | in 30 days.
20°6 25°5 16°59
20—25 mm. 21 27 19°93
| 23-4. | 27 10°73
(25 O77 7°53
rm | ) 26 29°5 9°41
25—30 mm. 5 265 38 3:95
| (27 312 10°85
40—45 mm. ; tae z ° a
|
45—50 mm. 46 | 46°5 ‘76
TaBLe LV.
Growth from July 31 to Sept. 7 (38 days). The Cloch, Gourock.
| Growth per cent. |
Group. Original Size. | Final Size. | in 30 days. |
16-4 219 | 2648 |
15—20 mm. 18 21 13°16 |
19°8 | 23'5 14:75 |
| |
35—40 mm. 391 43°6 | 9-08 |
Le f 42°5 43°5 1°85 |
aR Se ae 43°5 45°7 3°99
(46-2 AT 1:37
eee | 46°5 ATS 1-69
45—50 mm. < 46°8 Ar 1-01
1909. | OF THE LIMPET PATELLA VULGATA. 245
Tasue V.
Growth during July 1906 at Kames, Kyles ot Bute.
| Fl | | igeeete
| Group. Original Size. Final Size. | No. of days. | Bane
[o— 5mm. A | Oi (ibe. |
| | | | |
femora eer 1,6. 26) sea
| | 10 13 | 25 | 36°00
wom
| 12 | 125 31 | 4:03
| | 4
The observations recorded in Table I. amount to 42 for the
month of August.
The mean values calculated are given in Table VI.
TaBLeE VI.
Rate of Growth and Size.
Fe = Growth per cent. |
Group. No. | in30days. |
0— 5mm. 1 | 30:00
5—N0) x 2 16°84.
10—15 _,, 5 19°20
15—20 ,, 12 15°56
20—25 ,, 17 19°42
“25—30 ,, uit 6°89
30—35_,, 5 5°05
35—40_,, 5 4°77
40—45 ,, 6 2°71
45—50_,, 5 | 1:06
69
The percentages in the first two groups cannot be taken into
account since the numbers are so small. From 10 mm. on to
25 mm. there is a gradual and uniform falling-off in the rate ot
growth. In the 25-30 mm. group there is a sudden fall of the
vate to little more than half what it was at 20-25 mm., then
there is another gradual decrease up to the length of 50 mm.
There is a second slightly marked fall at a length of about 40 mm,
4. Seasonal Variation in Rate of Growth.
That the limpet grows more quickly in the warmer months of
the year is a fact which 1s clearly shown by the data in Table I.
In the first year-group for example, during the months of June,
July, and August, the monthly increment of growth is about
3 mm. During September the increment is 2°2 mm., during
246 MR. E. 8S. RUSSELL ON THE SHELL-GROWTH | Mar. 2,
October 1:1 mm., during November ‘6 mm.,and during December
‘T mm. ‘The percentage increase falls even more rapidly.
But it is in the second year-group (where there is little
variation in the rate of growth due to age) that the seasonal
variation is shown most clearly. During the first three months
of the year growth is practically at a standstill, then during the
summer a slow increase In size is manifested, but after October
growth almost ceases again. It should be remarked in passing
that the jump from 33-7 mm. at J uly 30th to 36 mm. at Sep-
tember 2nd does not signify that growth is specially active durmg
August. The increase is due to the introduction into the group
on September 2nd of the large shell No. 65 and to the reappearance
of No. 64.
The seasonal variation in the rate of growth may be proved in
another way. In Table VI. the percentage growth during the
month of August is tabulated for limpets of all sizes.
It is easy to calculate from these data the number of days
which a limpet would take to grow from 10 mm. to 15 mm., from
15 mm. to 20 mm. and so on, if it grew as quickly during the
other months of the year as it does during August. Using the
formula
OG 4b IP
oa LOO y sa
where O = original length, F = final length, P= percentage
increase per 30 days, and 2 = number of Tmo (of 30 days)
taken to grow from O to F, we obtain the following results :—
Time taken to grow Days. Sums.
‘hove 10) wawTeY, (HO) IS) WITN, 55545 Joaccncne one 69°26 69°26
15 Fe EO Rest ae ie ho Peele 59°71 128'97
20 pie ay ae pre BoC 57:18 18615
Ns acon’ Cilia teen mete at os 82°08 268°23
30 IS OV ieee aise ae Sore 93°87 362°10
35 pO aren ie ee vt Rae eee 100°77 462°87
40 sed eA edna re ee 132715 595°02
45 Sp OOMNae iitosacansedadace 30633 901°35
Suppose we start with a shell which is 10 mm. long on
January lst. If growth during the first two or three months is
as rapid as it is during August, the shells in the first year-group
should be 15 mm. long on March 10th, 20 mm. long on May 9th,
25 mm. long on July 5th. But actually, as is shown in Table I.,
they reach a length of 15 mm. only by the end of May, and
20 mm. some time towards the end of July. Hence growth during
late winter and early spring is slower than during autumn. The
same method might be applied to show that growth is again
retarded on the approach of winter, but one more application
of the method will suffice. If we take the second year-group
and start with a shell 30 mm. long on January Ist, it will, if it
erows at summer rates, reach a length of 35 mm. on April 4th,
1909. | OF THE LIMPET PATELLA VULGATA. DAT
40 mm. on July 14th, and 45 mm. on Nov. 23rd. But actually the
second year-group attains to a length of only 38 mm. even by
December 31st.
5. Rate of Growth and Reproduction.
We have seen that the variations in the rate of growth of the
limpet’s shell depend upon the age of the shell and upon the
temperature. It is natural to enquire whether the attainment
of sexual maturity has any marked effect wpon the rate of growth.
In other words, does arrival at a state of maturity coincide
with any marked drop in the rate of growth? It would seem
that it does, for sexual maturity is attained between 20 mm. and
25 mm., and there is a marked fall of the rate of growth at
25 mm. to half what it was at about 20 mm. Growth, however,
continues for long after a length of 25 mm. is reached, and ceases
only at 50-60 mm. The average full-grown size is about 45-
50 mm. The limpet shell therefore doubles its length after
sexual maturity has been reached. This fact recalls the law of
growth which Dr. Fulton has formulated for fish, that they
‘approximately double their size and increase their weight about
eight times after they have reached sexual maturity.” (4. p. 182.)
After the attainment of sexual maturity the shell becomes
much thicker and more solid in build. This increase in thickness
is no doubt correlated with its slower growth.
6. History of the Growing Limpet.
The facts detailed above enable us to form a mental picture of
the life-history of the limpet once it has settled down upon the
shore as a miniature adult.
It may come into existence at any time from July to January.
Its larval existence lasts probably only a week or two and it
settles down as a minute shell, one or two mm. in length.
Growth is rapid during the autumn. By January the shell is on
the average about 10 mm. long, and during January and February
it grows say 2mm. The rate of growth is then accelerated, and
growth continues actively throughout the summer. In August or
September the limpet, which is now 20-25 mm. in length, becomes
sexually mature. The rate of growth, which has hitherto been
anything from 12—20 per cent. per month, now falls abruptly on
the approach of winter and after the ripening of the gonad. By
the month of December the limpet is on the average 29 mm.
long. Very little increase of size is manifested during the early
months of the second year, but by April a slow growth sets in at
the rate of 5-7 per cent. per month, which falls off after the
summer is past to 1-2 per cent. and finally in the early winter
months ceases almost completely. During this year the gonad
has undergone profound changes, returning in the early spring to
a state almost of primary indifference and gradually ripening as
the summer advances. The gonad attains its full size in this
248 MR. E. 8S. RUSSELL ON THE SHELL-GROWTH [ Mar. 2,
second year, at the end of which the limpet is about 38 mm. long.
The sequence of events during the third year is probably very similar
to what it is in the second. Growth is even slower, and the
limpet may increase only 5 mm. in length. The limpet possibly
becomes full-grown at the end of this third year or at the end of
the fourth year—at a length of a little more than 45 mm.
Growth is almost at a standstill during the fourth year, and shells
of 48 mm. and upwards may exhibit for long periods no growth
at all, so that their age may be considerably more than four or
five years.
The distribution of lengths in a large random sample (Table IT.)
shows that the limpets are subject to a considerable degree of
elimination, the numbers in the first year-group (say 9-23-5 mm.
for the month of July) being much greater than the numbers for
the second year-group (say 24-38°5 mm.). The third and subse-
quent year-groups are not shown separately by a random sample
but merge into one another. The great number of shells 45—
53°5 mm. long which occur in the sample show that growth at
this size is practically stationary, for shells of this size represent
several different year-groups.
It is of interest to compare the life-history of the Tortoiseshell
Limpet, demea testudinalis, as related by Willcox (11). It shows
much analogy with that of Patella vulgata. Acmea testudinualis,
it should be mentioned, reaches a length of only some 20 mm.
The breeding-season in America lasts from the middle of April
to the end of July. Boutan (1) says that “ la ponte de /Acmée a
lieu en avril et en mai” on the French coast, and I have reason
to think that the same holds for the Acemea of the Clyde.
Willcox finds that the young dAcmea are 4-5 mm. long in
September and early October, and that they attain sexual
maturity under 10 mm., probably after one winter.
In Plate XXXII. the first two rows show stages of about fifteen
months’ growth, the second and third limpets in the second row
showing the size at which sexual maturity is reached. The first
two in the third row show the size of second year shells, the
second (38°5 mm.) being from two to two and a half years old.
The first limpet in the fourth row is probably a little more than
three years old, while the other two in the row are full-grown
limpets at least four to five years old. The two in the bottom
row are veterans, the last one (61°2 mm. in length) being the
largest limpet I have ever collected.
Il. CHANGES IN SHAPE DURING GROWTH.
About 1000 shells were collected at random during July 1903
and July 1904 at the Southend of Arran, and the ratios of the
various dimensions calculated for every shell.
Table VII. gives the mean ratios in each group; JL stands
for length, B for greatest width, H for height, Ab and Af are
explained a few lines further on
1909.] OF THE LIMPET PATELLA VULGATA. 249
TABLE VII.
Mean Ratios.
| L Ab L H
Group. No. ! B At: ‘ar B
| 5—10 mm. Ne 1:32 1:64 3°52 375
, OS 109 129 | 1:57 3°66 ta 352
| 150) 163 1:27 1:46 2:95 “431
| 20-25 . | 91 1:25 1:44 3°33 ‘375
(DRS) So VGH 1:23 1:36 Ss genes os
2035 68 121 1:34 PRey( ally 2, ED
35—40 ,, 81 119 131 265 | 449
| 4045. 115 ii 1:26 DS al ml
\ ZIG 0) 9 138 116 1:24 2°28 509
| 5055 ,, 122 115 1:22 2°16 532
| 55—60 ,, 25 115 77 2:28 504
As is eT in Table VII. very considerable changes take
place in the shape of the limpet-shell during its growth from
5 mm. to 60 mm. long.
L 5 E F
po Lhe shell becomes gradually broader in proportion to its
length, and this change is remarkably uniform from group to
group.
W pol’ grows higher in proportion to its length and in
proportion to its breadth. This change is on the whole a con-
tinuous one, but a reversal of its direction in both cases takes
place in the transition from the first to the second, from the
third to the fourth, and from the second last to the last groups.
Ab
IG
anterior margin (Af) becomes greater with great uniformity in
proportion oy the distance between the apex and the posterior
margin (Ab). It does not follow that the apex shifts backwards.
The changes in the position of the apex may be represented graphic-
i 5 : hi FF iN
ally if the successive values of the ratios Ht and a are drawn
to scale on a constant L. It appears from such figures that the
apex of the shell shifts gradually backwards during growth from
5 mm. to 25 mm., and then remains practically stationary till a
leneth of 55 mm. is reached, when it again shifts backwards.
Most of these changes can be directly inferred from the shape
of a typical fully-grown limpet-shell. A small limpet-shell has
the shape of a flattened excentrié cone, and the apex is in
such a young shell finely pointed and turned forwards. As
growth proceeds the sides of the cone do not grow straight
on, but begin to curve inwards slightly, so that by the time the
shell reaches a length of 50 mm. its outline as seen from the side
may be such as is shown in the Plate (last row). All good-sized
limpet- shells show this ‘ingrowth ” in the anterior and posterior
regions of the shell, and alge at the sides of the shell, for a similar
ee ingrowth ” takes place there also. This general “ingrowth ”
Proc. Zoou. Soc.—1909, No. XVII. eouial 77
—The distance in a straight line between the apex and the
250 MR. E. 8S. RUSSELL ON THE SHELL-GROWTH | Mar. 2,
5 < oo 1h :
accounts for the decrease of the ratio ul and the inerease of the
: : oa AUD Is
ratiop. The change in the ratio = is chiefly due to the same
>
cause, and especially to the fact, easily observed, that the
“inerowth” at the posterior margin is much greater in extent
than it is at the anterior margin, whence it arises that Af becomes
L
larger in proportion to Ab. In the three cases in which a
increases and q decreases during growth from one group to the
next, one must suppose that an “ outgrowth” has taken place,
caused probably by the formation of very large projecting ribs, or
else that the apex of the shell has been ab ‘raded. The latter
explanation is probably the true one for the 55-60 mm. shells,
for such old shells are usually very much worn and eroded,
especially towards the apex.
The change in the ratio a may possibly be due also in part
to the unequal rates of growth of the anterior and posterior
margins. There is some reason to think that the anterior margin
grows a little quicker than the posterior. If one examine a
well-preserved “smooth” shell (see 9. p. 868) from above, one
sees numerous fine ribs radiating from the apex. Now it very
often happens that many of the lateral ribs curve slightly
forward, while the anterior and posterior ribs run out in a
straight line from the central apex. This shows that during
growth the mantle (or the whole limpet perhaps) gets shitted
forward a little relatively to the apex, and that accordingly the
growth of the anterior margin becomes somewhat greater im
proportion to that of the posterior margin. The ribs of the shell
end in little projections of the rim, and to these projections of
the rim correspond little projections of the edge of the mantle,
hence from a curving forward of the ribs one can infer a
movement forward of the mantle relatively to the shell-apex.
Tn addition to the radiating ribs there are visible on unworn
shells numerous lines of growth. These are often very distinct,
especially at about the level corresponding to lengths of 15 mm.
and 30 mm. The shell lying between these two levels is probably
added during the first summer and autumn’s growth. It is,
however, hardly possible to recognise annual lines of growth on
the shell other than these two.
Shells of 40 mm. and more are usually very much worn on
the upper part, from a level of 30 mm. upwards.
Besides the changes in shape during growth there is also a
marked change in the build of the shell. It becomes very much
thicker and heavier. This increase in thickness (out of proportion
to the increase in length) becomes noticeable from a length of
about 25-30 mm. onwards, 7. e. in one year old shells and older
ones. The rim becomes considerably thickened, and a great deal
of shell-material is laid down inside the apex, which soon becomes
solid to quite a considerable depth. In a limpet of 60 mm. the
thickness of the shell at the apex was so great as 10 mm.
1909. | OF THE LIMPET PATELLA VULGATA. PHYA
Throughout all these changes there persists a great uniformity.
The relation that L>B>Ab>Af>H had no exceptions in the
one thousand and three shells measured, except that occasionally
in small shells B equalled Ab. ‘The apex is wlways nearer to the
anterior margin than to the posterior margin. The greatest
breadth is invariably behind the apex, and ‘usually behind the
middle point of the shell, so that the outline of the margin is not
an ellipse. Some results published by Malard (7) are “obtained
upon the assumption that the outline of the shell-margin may he
treated as an ellipse. The index of ellipticity is calculated for
several subspecies of Patella, and also an index of height. It
does not appear that any allowance v as made for the differences
in the ratio of length to breadth and ob each to height, which are
found in shells of different lengths. For a form collected at.
Barfleur the index of ellipticity was 6908; the curve of fr equency
was asymmetric and of Type J. Malard’ s inference that in this
case selection was more pronounced in one divection than the
other is quite unjustified.
The question naturally arises, are these changes in shape
simply due to “laws of growth,” or are they brought about
by selection? It seems fairly certain that the changes in the
‘atios of the various dimensions of the shell are not due to the
action of a process of natural selection, and that for two reasons :
first, the changes are perfectly continuous and gradual from
stage to stage ; “second, they are exhibited by every shell without
exception. If the changes were due in any marked degree to the
action of natural selection, a process of differential destruction
must have been active at all stages, and one would expect to find
dead shells which did not exhibit in their shape the changes
which all good-sized limpets show. But such shells ave not to be
met with.
It can indeed be proved directly that the changes in shape
may take place without the action of natural selection For
instance, 76 shells of 40-45 mm. length were collected at the
Cloch, Gourock, and the mean breadth at a length of 40-42°5 mm.
was calculated. The value obtained was 35°1448 mm., and his
ratio of length to breadth was 1174. The mean breadth at +
length of 15-17-5 mm. was also calculated from measurements of
50 to 60 of these shells, advantage being taken of the lines of
growth visible on the shell. (A few shells 38-40 mm. in length,
collected at the same time as the 40-45 mm. set, were measured
in order to make up ee number to 60.) The ratio of length to
breadth at this length was 1:26. The ratio of length to breadth
in 60 shells 15- 17-5 mm. in length collected at tite same place
and time as the large shells was 1:24. Now within the group
composed of the 60 large shells natural selection has not acted,
for natural selection Le by elimination, and there has been no
elimination within the group. It is true that selection may have
been at work before the ler vgth of 15-17°5 mm. was reached, and
that these shells may r epresent a selected sample of the population
below 15-17°5 mm. It is true also that natural selection me ay
Lit
252 THE SHELL-GROWTH OF THE LIMPET PATELLA VULGATA. [ Mar. 2,
have been at work on other shells 15-42°5 mm. in length—-that
possibility always remains. But for this group the action of
natural selection between the lengths of 15 mm. and 42°5 mm.
is excluded. Hence all the changes taking place during this
period of growth are independent of selection. Now the values
obtained for the ratio of length to breadth approximate very
closely to those obtained for Groups 15-20 mm. and 40-45 min.
of the large series (Table VII.), and though the value for the
breadth of the 40—42°5 shells at 15-17-5 mm. long (1°26) falls some-
what below the value of the ratio in Group 15-20 mm. (1°27),
and might lead one to think that these shells were a selected lot,
yet the value of the ratio for the 60 shells 15-17°5 mm. in length
is even more divergent, being only 1:24. The differences are
clearly due to the difference in environment, the series of which
the ratios are tabulated in Table VII. having been collected in the
Southend of Arran. One objection may be made to these results,
and that is that shells of 40-45 mm. are usually very much
abraded towards the apex, and consequently the measurements
made on them at a length of 15-17°-5 mm. cannot be depended
upon. This objection has some weight, and to corroborate my
results I have made measurements of 100 “smooth” shells
25-35 mm. in length (75 of them being 25-30 mm. long), which
were carefully selected from over two hundred very kindly collected
for me by Mr. David Baird, M.A., from the shore where my
original set of 1000 had been gathered. The shells selected were
those which had suftered practically nothing from weathering,
and I am certain that the results they give are not appreciably
influenced by the effects of abrasion and erosion. The ratio ata
length of 15-17:5 mm. on these shells was 1°22, and at a length
of 24-26°5 mm.it was 1:19. The change in this ratio may quite
certainly be set down toa “law of growth” and not to selection.
This change comes about in approximately four months’ time.
Summary.
The breeding-season extends from July to January.
Sexual maturity is reached at a length of 20-25 mm.
An average size for a limpet of the last season’s brood in
January or Februaryis 10 mm. At the end of the first year
it may be 29 mm. long. Probable sizes at the end of the
second and subsequent years are 38 mm., 44 mm., 48 mm.,
53 mm. Shells over 50 mm. may be considerably more than
five years old.
(4) Sexual maturity is reached in the first year, and when the
limpet is only half-grown.
(5) The rate of growth decreases with age and maturity. It is
slower during the colder months of the year.
(6) Considerable changes take place in the ratios of the shell-
dimensions during growth.
(7) These changes are probably in large part the expression of
“laws of growth,” and not due to natural selection.
Fa a i
(SG) (SS) =
VSS
Ten ApS WO) dell DOOKU,
West,Newman lith.
DEVELOPMENT OF AGRIONID DRAGONFUIES.
P.Zo8: 1909 PY OOGIN:
West,Newman lith.
DEVELOPMENT OF AGRIONID DRAGONFLIES.
i)
MS)
1909.] ON THE LIFE-HISTORY OF THE AGRIONID DRAGONFLY. 5
REFERENCES.
(1) Bouran.—C. R. Acad. Sci. exxvi. (1898) pp. 1887-89.
(2) Davis, J. R. A., & Fururs, H. J.—“ Patella.” Liverpool
Marine Biological Committee Memoirs. Memoir x. (1903).
(3) Fiscuer, P.—Manuel de Conchyliologie. Paris, 1887.
(4) Furron, T. Wemyss.—24th Ann. Rep. Fish. Board Scot.
(1905) pp. 179-274.
(5) Gemnity, J. F—Anat. Anz. xii. (1896) pp. 392-4.
(6) Lesert et Ropin.—Ann. Sci. Nat., Zool. (8) v. (1846).
Ppa lol 2.
(7) Mauarp, A. E.—Bull. Mus. d’Hist. nat. Paris, ix. (1903)
pp. 270-4.
(8) Parren, W.—Zool. Anz. viii. (1885) pp. 236-7.
(9) Russetn, E. $.—Proc. Zool. Soc. 1907, pp. 856-70.
(10) Semper, K.—“ Animal Life.” London, 1881.
(11) Witucox, M. A.—Amer. Nat. xxxix. (1905) pp. 325-33.
(12) Witson, E. B.—Journ. Exper. Zool. i. (1904) pp. 197-268.
EXPLANATION OF PLATE XXXII.
SHELLs oF Patella vulgata L.
The actual lengths of the shells are as follows —
Ist Row.—7 mm., 89 mm., 10 mm., 11 mm., 13°5 mm., 15°4 mm., 17°2 mm.
2nd Row.—20°8 mm., 23 mm. (sexually mature), 266 mm. (about 12 months
old), 30 mm. (15 months).
3rd Row.—33'3 mm., 38°5 mm. (over 2 years), 40°7 mm.
4th Row.—44'1 mm. (over 3 years), 47°3 mm. (about 4 years), 51°2 mm. (about
5 years).
5th Row.—53’5 mm., 61°2 mm. These two limpets are at least six years old, and
may be more.
3. The Life-History of the Agrionid Dragonfly.
By Frank Barrour-Browne, M.A. (Oxon.), F.R.S.E., F.Z.8.
[Received February 5, 1909. |
(Plates XX XIII. & XX XIV.*)
INTRODUCTION.
So far as I have been able to find, the life-history of the
Dragonfly has not, up to the present, been studied in detail by
anyone. Various facts have been noted by different observers,
but no one has apparently attempted to hatch the eggs and follow
out the changes which take place during the larval or nymphal
condition.
The general facts in the life-history are well known. The
perfect insect lays its eggs in the water; after an unknown
period there emerges from the egg a sub-aquatic creature known
as a nymph, which grows, casting its skin at unknown intervals,
* For explanation of the Plates see p. 285.
254 MR. F. BALFOUR-BROWNE ON THE [ Mar. 2,
and which finally crawls out of the water and dries itself in
preparation for the final ecdysis.
In this paper I propose to describe the details of the deve-
lopment so far as I have been able to ascertain them ; and, while
so doing, I shall incorporate a few casual observations on the
anatomy which have been made in the course of the work and from
watching the living creatures themselves. Although incomplete,
these observations are, I believe, accurate, and may prove of
some use to those interested in the comparative anatomy of the
Insecta.
I commenced studying the Dragonfly in 1903, at the Sutton
Broad Laboratory; but it was not until July 1906 that I seriously
undertook the investigation of the life-history. From the litera-
ture on the Odonata I was able to obtain but little assistance.
Packard (1866) published a paper on the development of the
Dragonfly Diplax; but that paper deals for the most part with
embryology, and the illustration of the newly-hatched larva—
which is reproduced by Dr. Sharp, (1895, p. 419),—although truly
representing the first stage of the true nymph, does not represent
the creature as it comes out of the egg. Brandt, Dufour, Amans,
Gilson, and various others who have worked at the Odonata have
dealt with their anatomy and physiology, but not with their life-
history; and W. J. Lucas (1900), in his book on the British
Dragonflies, only refers in a general way to the life-history. He
also makes the mistake of regarding the first stage of the true
nymph of Sympetrum str iolatwm as the form lel emerges from
the egg.
It was necessary therefore to start de novo, and for this
purpose I endeavoured to collect some eges from the dykes in
the neighbourhood of the Laboratory. The females of all the
Zygopteride, or thin-bodied dragonflies, like those of the
AMschnide but unlike those of other Anisopterids such as Libel-
lulide, Gomphidee, &c., are possessed of an ovipositor and lay
their eggs in the tissues of floating aquatic plants. ‘They settle
on a floating leaf of Hydrocharis, Potamogeton, Anacharis, &c.,
and dip the end of the abdomen into the water, often immersing
the whole of it. According to some books, the females actually
go under water at times when ovipositing; but I have never
seen this. In the case of those plants with broad floating leaves,
the apex of the abdomen is curved under the leaf so ‘that the
ovipositor, which is on the ventral side of the 9th abdominal
segment, is brought into contact with the lower surface. The
ovipositor consists of two equal lobes placed side by side, each
with a sharp point projecting backward. The two halves of the
ovipositor work together and pierce a small hole in the leaf-
tissue. They then separate slightly from each other, and
between them a single elongated egg passes down and becomes
wedged in the hole in the leaf. Only one egg is buried in each
hole. Some discrimination is exercised as to the piercing of the
plant-tissues, as the ovipositor moves about over the surface
1909. | LIFE-HISTORY OF THE AGRIONID DRAGONFLY. 255
before actually entering the leaf. The eggs are easily found in
the thin leaves by holding these up to the light.
East Norfolk is rich in “dragonflies * , not only in species but in
individuals. A great number of females of Agrion pulchellum
Lind., /schnura elegans (Lind.), Hrythromma naias (Hansen),
Lestes sponsa (Hansen), and Hnallagma cyathigerum (Charp.)
were ovipositing at Sutton in the end of July. Eggs were
therefore not difficult to find; but it was a somewhat laborious
task to collect a large number, and, apart from this, it was
impossible to know to which species any egg belonged or how
long it had been laid. I therefore constructed a “large cage,
15" x 12" x 12", consisting of a wire frame covered with muslin,
and this was placed over a large photographie developing-dish
half filled with water. On the water I floated a few clean young
leaves of the frog-bit (ydrocharis morsus-rane L.) cut off from
the plant, having first examined them carefully to make sure they
contained no dragonfly egos. I then captured six pairs of Agrion
pulchellum Lind. and put them in the cage. It is extraordinary
how dependent upon the direct sunlight is the vitality of these
insects. On dull days, or when the cage was in the shade, the
prisoners were in what appeared to bea torpid condition, clinging
to the muslin sides of the cage and not moving for hours at a time ;
but on a bright day, or as soon as the cage was put into the sun-
light, the scene became one of extraordinary activity: the males
fighting one another for the females, and the combatants con-
stantly falling into the water, from which, however, they always
succeeded in escaping.
The peculiarity of the male reproductive organs—the vasa
deferentia opening on the sternite of the 9th abdominal segment,
while the organ of intromission is situated on that of the 2nd
seement,—and the consequent peculiar method of transference of
the spermatozoa to the female are apparently unique in the
Odonata. I have never seen the actual transfer of the sperm to
the sperm-sacs of the male, although in some individuals which
have emerged in captivity I have noticed the tendency to curl the ©
abdomen asif making attempts to bring the 9th and 2nd abdominal
sternites into contact. Whether the sperm is transferred to the
sacs once and for all, or whether the supply in the sacs is
replenished from time to time, has apparently not been observed ;
but the female, at intervals in the process of oviposition, obtains
further supplies of sperm from the male.
The male is furnished with claspers at the apex of the abdomen,
by which he grips the notum of the prothorax of the female; and
the pair fly about attached in this manner. The adaptation of
these parts to one another is so close that only the claspers of the
male of its own species can hold the prothorax of a female. Thus
one very definite means of separating the females of A. pulchellum
* See ‘ Victoria County History: Norfolk, vol. i. (1901). Also, for Agrionide,
«A Bionomical Investigation cf the Norfolk Broads,’ Trans. Nortolk & Norwich Nat.
Soe. vol. vii. (1904).
256 MR. F, BALFOUR-BROWNE ON THE { Mar. 2,
and A. puella, which very closely resemble each other, is the form
of the notum of the prothorax, and I give drawings of this part of
both these species. It will be seen that the males also differ some-
what in this respect, but to a less extent than the females.
Hither while a pair are flying about attached per collem, or
while so resting, the female curves her abdomen underneath her
in such a way as to bring the underside of her 9th segment into
contact with that of the 2nd abdominal segment of the male ; and
the parts are so constructed that a firm attachment can take place,
and the spermatozoa are transferred from the sperm-sacs of the
male to the oviducts of the female.
After a contact of shorter or longer duration, from a few seconds
to a minute or more, the female once more straightens herself out
and sooner or later oviposition commences. This is most commonly
done while the male is still holding the female per collem, but,
though I believe less commonly, the female will oviposit without
being held bythe male. The male remains entirely passive during
oviposition, apparently obeying the will of the female. When she
flies, he also flies; but she directs, or seems to direct, the course.
She seeks the aquatic vegetation and deposits her eggs as already
explained. The male anenmernille rests either with his legs on the
leaf or more commonly standing erect in the air, his body rigid,
his legs folded, and himself supported only by the claspers attached
to the prothorax of the female.
I kept the prisoners in the cage for about a week, feeding them
on flies and other insects captured with the sweeping-net. The
prey crawled about on the muslin, and the dragonflies, flying about
in the cage, picked them off and either ate them while still flying
or after settling onthe sides. A large number of eggs were mean-
while deposited in the frog-bit leaves, besides very many which I
found in the bottom of the dish. The eggs were laid during the
first four days of August, and I placed the frog-bit leaves in
tumblers of clean water and also collected a number of the loose
eges and dealt similarly with them.
After about a fortnight the leaves showed signs of decay ; and
towards the end of August nothing of them remained but a mass
of decayed vegetation at the bottom of each of the tumblers, and
my hopes of seeing any nymphs from the eggs vanished. How-
ever, on the 28th ‘of the month numbers of nymphs appeared on
the top of the mass of decay—almost four weeks after the depo-
sition of the eggs. Although the majority of the eggs hatched at
one time, nymphs were appearing for a fortnight or three weeks
after that ; and the same fact was noticed with the eggs of /schnura
elegans on another occasion.
The eggs which were dropped in the bottom of the dish never
developed at all, and were all decomposing within 10 days. I
have found that the eggs of the water-beetle Dytiscus also failed
to hatch when cast loose instead of being buried in plant-tissues,
as they normally are; and Miall (1895, p. 40) mentions that
Lyonnet also found that loose eggs of Dytisews never hatched.
1909.] LIFE-HISTORY OF THE AGRIONID DRAGONFLY. 207
It has occurred to me that this failure of loose eggs is due to
asphyxiation, as possibly the eggs buried in the plant-tissues
obtain a certain amount of air from the intercellular spaces
of the plant; but if this is the case, it is somewhat surprising
that the decaying leaves of the frog-bit retamed sufficient air to
supply a fair number of eggs.
At the time when the nymphs appeared I did not observe the
emergence of the nymph from the egg; but I have since had
the opportunity of watching nymphs of /schnura elegans hatch,
and I have no doubt, from the similarity in all the later stages,
that the peculiar phenomena observed in the case of J. Asseuus
oceur in all the Agrionide, and, from the fact that I have
observed some of the same phenomena in one of the Aniso-
pterids, Sympetrum striolatwm, they are evidently common to all
the Odonata.
The dragonfly, both as a nymph and as an imago, requires
living animals as food, and at the commencement of the work the
difficulty was to find food small enough for the nymphs, which
were but little over 1 mm.in body length. An infusion of horse-
dung in water solved this difficulty, as in a few days it swarmed
with life, one jar in particular constantly producing, for five or
six months, a rich supply of Paramecia, which the nymphs ate
readily. As the nymphs grew the Paramecia were replaced by
daphnids, and it was surprising what large specimens the nymphs
caught and demolished. Daphnids served as food all through the
nymph stage after the protozoa became insufficient.
~ The nymphs proved exceedingly easy to rear. A number of
them, as soon as they appeared, were sorted out and measured,
and each was then placed in a tumbler of water by itself, a small
piece of Anacharis being put in to give it foothold and also to
keep the water fresh. Hach tumbler was then labelled, and
a record kept of the life of its occupant. The water in the
tumblers was only very occasionally changed, and only when
the glass became so thickly coated with algz that I could not
see the inmate. For the first two months of their hfe the
nymphs were examined and measured under the microscope at
least twice a week—for the first week or two, every other day ;
but such frequent interference with their existence proved
unnecessary, and latterly the examination and measurement
was much less frequent, being almost entirely confined to the
times of moulting.
Although I ine only followed Agrion pulchellum and /schnura
elegans right through from the egg to the imago, I have also
kept through many stages nymphs of Pyrrhosoma nymphula
and Hr; yihromnus naias, and from my observations I think there
is no doubt that, so far as the general facts go, what is true for
one of these species is true for all the others.
In this paper I am treating the subject from two points of
view. First, I have detailed the phenomena observed in relation
to the stages in the life-history in which they occurred, the stages
258 MR. F. BALFOUR-BROWNE ON THE [ Mar. 2,
being marked by the successive moults. I have then gone into
some detail with regard to the development of the form of the
nymph—that is, morphologically rather than anatomically; and
in this part of the subject one or two interesting facts as to the
manner of growth of the external parts have been observed.
This detail has brought to light a certain want of agreement
between the stages as marked by the moults and the stages in the
development of these external parts; and I have therefore, in
conclusion, reviewed the question from the point of view of the
significance of the moult. This latter part of the paper is very
incomplete, since, although a number of facts on the subject of
the moult in insects are known, there is insufficient material at
present upon which to base any really satisfactory theory. How-
ever, this is merely forestalling the conclusion.
THe PRoNYMPH.
The Agrionid egg is an elongated body, rounded at one end and
bluntly pointed at the other, the point being a small pedicel. In
Agrion pulchellumand Ischnura elegans its length is about 1°3mm.,
but the size no doubt varies in the different species. The egg is
inserted, as already explained, in the soft tissues of aquatic plants,
the pedicel end being the last to leave the oviduct and being
therefore nearer to the surface of the leaf in which the egg is
deposited.
The period of incubation varies somewhat in length according
to temperature, apart from the fact that all the eggs of a batch do
not hatch on the same day. Eggs laid inthe beginning of August
took just less than four weeks Tatlone the rly of fhen hatched,
while eggs laid later in the year did not hatch for more than five
weeks.
For some time previous to hatching the position of the nymph
within the shell is easily made out. The head lies at the pedicel
end of the egg, the antenne, first maxille, and labium being
directed backward on the ventral side of it. The first and second
pairs of legs le straight out along the ventral side of the abdomen,
while the third pair are bent on themselves at about the junction
of the tibiz and tarsi, the latter lying forward below the tibie.
The posterior abdominal segments curve round in the shell
ventrally, the 10th occupying the rounded end of the shell. The
caudal lamellee lie forward along the ventral side, between the legs
and the shell. :
For at least three days before the egg is ready to hatch the heart
can be seen beating, at first slowly but getting faster as the time
approaches for the emergence of the pronymph. The posterior
end of the heart is the part which develops and pulsates first, and
although in the free nymph this is in the 9th abdominal segment,
in the egg it appears to lie in the 10th segment.
The first sign of the approaching emergence of the pronymph
is the cracking of the shell immediately beneath the pedicel. At
1909. | LIFE-HISTORY OF THE AGRIONID DRAGONFLY. 209
this time the heart is beating vigorously, at about 100 pulsations
per minute. The pedicel is then seen to be slightly removed from
the egg, owing to the bulging out from the egg of an extremely
fine membrane which is evidently extended with fluid. This
membrane continues to expand until there is a gap of about
-15 mm. or even more between the pedicel and the rest of the
shell. It is noticeable also that the pedicel is not carried up in a
line with the axis of the egg, but rather dorsally to the axis, and
although this process was only watched in two or three cases
I believe that the direction of expansion of the membrane is
perfectly definite.
This carrying up of the pedicel and the formation of the vesicle
occupies one or more hours, and except for some occasional shght
writhing movements of the gut and of the abdominal segments
within the shell no other change is noticeable during that time.
The vesicle is, in fact, formed many hours before the pronymph
begins to emerge. Some time after the formation of the vesicle
a pulsating organ appears in the head in the region of the mouth ;
and one or two hours after this organ begins to pulsate, the head
of the pronymph begins to expand into the vesicle. It takes the
shape of the passage, losing all semblance of a head, and slowly
fills the vesicle until it once more touches the pedicel. All this
time the pulsating organ continues to beat at about 28 beats per
minute.
As the head begins to enter the vesicle the walls of the latter
bulge slightly, showing that they are extended by fluid which is
being compressed by the head. A]most as soon as the head com-
pletely occupies the vesicle the membrane bursts along the dorsal
edge of the pedicel and the enclosed pronymph slowly comes out.
Just previous to this body-movements are more noticeable, and the
tenth abdominal segment moves round so as to be in line with the
others; but until the membrane bursts, no part of the shell is
unoccupied.
The pronymph appears to flow out of the shell at the start, but
almost immediately it exerts itself and wriggles from side to side.
It comes out in exactly the position in which it has lain in the
shell; the lamelle, bent under the body, coming out points first.
The lamelle straighten out more or less once the larva is clear of
the shell, but the legs and mouth-parts remain outstretchea and
directed backwards.
From the moment of its escape the pronymph—a helpless
cripple—shows active writhing movements, and within two or
three minutes the skin on the back of the thorax splits along the
median line and a moult takes place. The posterior abdominal
segments contract and expand and gradually creep up inside the
pronymph skin, drawing the lamelle out of their sheaths. As the
abdominal segments press forward, the thorax becomes arched up
and the head bent down. ‘The pressure in the dorsal region of the
thorax ruptures the skin along the median line, and the thorax
rises up through the split. In doing soit ruptures the skin further
260 MR. F, BALFOUR-BROWNE ON THE [ Mar. 2,
forward, and the head with the antenne and mouth-parts is drawn
backward and upward out of the old skin and all these parts at once
assume their normal position—the antenne pointing forwards, the
labium doubling on itself and taking up its position below the
mouth, the maxillz likewise bending forwards and coming to le
on either side of the mouth immediately below the jaws. Further
writhing of the abdominal segments forces the body still further
out of the skin, and the legs are withdrawn from their sheaths.
Once the legs are free, a final struggle clears the nymph of the
pronymph skin. This latter is exceedingly thin and transparent,
and almost invisible even under the microscope except for the fact
that on each side of it is an irregular band of backwardly-turned
projecting points. These lie scattered on either side of the thoracic
and abdominal segments, but there is none on the head. These
points are no doubt of use in assisting the pronymph to escape
from the egg.
Owing to scarcity of material after the discovery of the above
details of the pronymph stage, I was able to observe the emerg-
ence from the egg only in two or three cases, and I had not
therefore opportunity of making many observations on the
pulsating organ already referred to. Several points were, how-
ever, noted. The organ is not far from the position of the
mouth ; it contracts from in front backwards, apparently driving
blood, since the corpuscles were visible, posteriorly, and the
pulsation and direction of the flow were quite distinct. The
drive was towards the dorsal surface near the posterior margin of
the head, and a biood-vessel in that region (the aorta?) could be
seen showing slight bulging each time the liquid was forced to it.
Blood could also be seen passing from the anterior part of the
head back to the pulsating organ; but at times this flow ceased
and the corpuscles appeared to oscillate backwards and forwards
at each beat of the organ, just as they may often be seen to do in
the ordinary blood-spaces of the body.
I could find no trace of a pulsating organ in the pronymph
after it had escaped from the egg; but this might have been
due to the fact that the creature was in constant movement
for the two or three minutes of its existence. There is
apparently no such organ in the nymph. I it only appears,
as L believe, after the formation of the vesicle and disappears
or ceases to pulsate as soon as the pronymph escapes from
the egg, the obvious inference is that it has something to do with
the emergence of the pronymph, and is, in fact, an embryonic
organ. It is interesting in this connection that Wheeler (1893)
has described an embryonic ‘ subeesophageal organ” as occurring
in insects, of which he says the cells somewhat resemble those of
the fat-body. He places it in the tritocerebral segment of the
head, and says that it disintegrates in the larva.
The function of the pulsating organ in the head of the Agrionid
embryo seems to be to absorb the fluid in the vesicle, so as to
allow the head to pass into it; but how it can do so is at present.
1909. | LIFE-HISTORY OF THE AGRIONID DRAGONFLY. 261
amystery. In the embryo of the water-beetle (Hydrobius fuscipes
L.) I find that there is a cephalic pulsating organ which becomes
active a few hours before the escape of the embryo from the egg,
and that during its activity the embryo enlarges so as to completely
fill the egg. There is, however, no vesicle formed, and in that
case I am almost certain the pulsating organ is not a special
structure but merely a pharyngeal pump, which is found in the
larva after its emergence and is similar to that in all suctorial
insects. In the Agrionid nymph, however, there is no suctorial
apparatus ; apart from which, the blood-corpuscles visible in the
pulsating organ oppose the idea that it is part of the alimentary
canal.
Civeulatory pumps, accessory to the heart, have already been
described in insects, but so far, I believe, they have only been
observed in imagines and ase and not as embryonic organs.
Behn (1835) first discovered them in the legs of various Hemipter a,
and Selvatico (1887), Pawlowa (1895), and others have described
them in the heads of certain Lepidoptera and Orthoptera.
Selvatico describes an enlargement at the anterior end of the
aorta—the ‘ frontal sac,” a description which might possibly suit
the organ in the Agrionid embryo.
The reversal of the blood-stream observed in the embryo is also
not unique*, as Bataillon (18935) observed the same phenomenon
in the pupa of the silkworm.
With regard to the formation of the vesicle, it is possible that
it is due to the displacement of the fluid which presumably
surrounds the embryo within the shell, or possibly the fluid is
extruded from the embryo to be later reabsorbed.
Much, however, remains to be discovered with regard to these
phenomena i in the egg.
THe Nymex. Stage 1.
The nymph which escapes from the pronymph skin is no longer
than the pronymph, in fact the total length—not including the
caudal lamellee—is just about that of the egg from which it came.
It consists of a head, 3 thoracic and 10 abdominal segments, and
is an excellent example of the campodeiform larva. The head
bears two antenne, each of which consists of 3 segments, two
basal ones, which are of characteristic form and except in size do
not show any change during the life of the nymph, and a long
bluntly pointed, slightly ineurved apical one. Not far from its
apex, and on its outer side, this apical segment bears a very small
spur. The length of this segment is equal to about 70 per cent.
of the whole antenna.
The mouth-parts consist of a pair of jaws of the usual
* I have recently observed a reversal of the blood-stream in the heart of a well-
grown nymph, and I imagine the blood was also passing into the last chamber of
the heart by the last pair of ostea. It was escaping by the ostea in the 8th abdominal
segment.
262 MR. F, BALFOUR-BROWNE ON THE [ Mar. 2,
“‘ mandibulate ” type, a pair of maxille, and a labium of very special
form usually known as “the mask.” It is in shape more or less
an isosceles triangle, the apex being turned posteriorly and lying,
when in a state of rest, underneath the first thoracic segment.
The base of the triangle is presented forwards and at each of the
anterior corners is attached an appendage representing a labial
palpus, which again is rather triangular in shape, being flattened
above and below and having three sides, one of which is anterior.
At its outer anterior extremity each palp bears a movable claw,
and along its anterior border each palp is beset with four or five
teeth, the innermost being much the largest and slightly curved
inwards. ‘The inner border of each palp is serrated, the teeth
being small and curved slightly backward. The anterior border
of the labium itself is also slightly serrated *, and when the mask
is not in use the palpi lie against this border in such a way that
the anterior edges of the two interlock, the movable claws over-
lapping. On the upper side of each palpus, attached near its
outer border and not far behind the movable claw, is a single
long incurved hair or bristle.
The thoracic segments bear each a pair of legs, but at this stage
there is no trace of the wings nor are the pleural ridges present.
Except for the possession of the legs there is nothing in out-
ward appearance to distinguish the thoracic from the abdominal
segments. These latter are also very similar to one another,
except that the body tapers slightly towards the posterior end.
The tenth segment bears the three caudal lamelle, one median
situated above the anus and two laterals, one on either side of
the anus. The two main trachee, which pass along each side of
the thorax and abdomen, curve inward and upward in the tenth
segment and run out side by side in the middle lamella, At the
points where they curve upward a single branch continues as a
fine tube into the lateral lamella of its own side. The lamelle at
this stage are tubular, perhaps slightly triangular in transverse
section, tapering to their apices and ending in a fine long hair ;
they bear along two or three lines a series of small hairs and in
Agrion pulchellum they contain two patches of pigment, one near
the apex and one about halfway from the base.
The transparency of the nymph at this stage enables one to see
fairly well the internal organs. The gut consists of a pharynx
leading to an esophagus which opens out into a large thin-walled
sac—which is apparently contractile—occupying the thoracic
segments. At the posterior end of this sac there is a constriction,
and immediately behind this is another large thin-walled sac
which occupies most of the first six abdominal segments. This
sac—the mid gut—is constricted posteriorly and opens into a small
globular contractile portion of the intestine, in the anterior end
of which open two Malpighian tubules, one on either side and
* The serration does not appear on either the palpi or the labium until a later
stage. Only faint traces of it are visible at this stage.
1909.] LIFE-HISTORY OF THE AGRIONID DRAGONFLY, 263,
shghtly dorsally. The rectum is the continuation of the small
intestine and occupies almost the whole of the last four segments
of the body. The small intestine at one time appears globular
and at another as a thick cord of tissue, the contractions appa-
rently commencing anteriorly and passing backwards. A. con-
traction begun in the small intestine continues through the
rectum, which, like the former, is sometimes a large sac and some-
times a thick cord. With regard to the number of Malphigian
tubules, Wheeler (1891-3) suggests that the embryonic number
in the Odonata “will probably be found to be 4 or 6.” There
are, however, only two in Agrion pulchellum at this stage.
The heart consists of nine compartments and lies in the
abdominal segments 1 to 9. Hach compartment is separated from
the next bya pair of very distinct valves, which liein the posterior
region of each abdominal segment. Immediately in front of each
pair of valves is a pair of ostiain the lateral walls of the heart, so
arranged that when the valves are open between two compart-
ments the ostia are closed. The heart opens anteriorly into the
dorsal vessel, or aorta, which has only a slight contractile power,
and this vessel runs forward into the anterior region of the head,
where the blood-stream divides into two and passes downwards in
front of the head.
There is at this stage no circulation in the antenne or in the
limbs. The blood returns through the body by various lacunz.
The two large lateral sinuses, in which at later stages the blood-
flow is easily seen, are but little developed at present. The
corpuscles, which are large more or less oat-shaped bodies, can be
seen passing backwards along the “ pericardium,’ and a few can
be seen every now and again making their way up from the
lateral sinuses and after much oscillation entering the heart by
some of the anterior ostia. The great majority of the corpuscles,
however, re-enter the heart by the two large ostia situated at the
posterior end of the 9th segment. Between these two ostia is a
strong ligament passing back into the 10th segment.
The ventral nervous system, so far as it is visible in the living
nymph, consists of a large (double) ganglion in each of the
abderminal segments 1 to 7, and between the ganglia the cord
is double, the two parts being quite separate. Behind segment 7
these two cords pass backwards and break up into finer nerves.
There is no sign at this stage of any sexual armature. This
first appears, I believe, about the 7th stage, but I have made very
few observations on this point.
Stage 2.
At the first moult of the true nymph certain changes are
visible, showing an advance in organization. . The antenne are
now composed of 4 segments, the apical one having divided
transversely into two of almost equal length. The labial palpi
now each bear two long stiff incurved hairs, the second hair having
264 MR. F, BALFOUR-BROWNE ON THE [ Mar. 2,
appeared below the first at the same distance from the external
edge of the palp and much nearer the base.
The lamelle show no change in shape, but the two lateral ones
now possess two tracheal tubes, the main branch as its enters the
base having given off a smaller branch which runs up the lamelle
parallel with the main one.
In some specimens at this stage circulation of the blood is
visible in the basal segments of the third pair of legs, but, as a
rule, there appeared to be no difference in this respect between
this stage and the previous one.
Stage 3.
The antenne now consist of 5 segments, the third from the
base having divided, cutting off about one-third of its length at the
distal end to produce the new segment.
No change takes place in the labial palpi, but one pair of long
stiff hairs has appeared on the upper surface of the labium itself.
These hairs are situated nearer the anterior than the posterior
end and are on each side of the median line fairly wide apart.
They are directed forwards and, when the labium is at rest below
the head, are immediately behind the mouth of the nymph.
The lamellae are slightly wider vertically than before, but 1¢ will
be necessary to deal with their development separately, as their
changes apparently correspond with the rate of growth of the
nymph rather than with the stage in its life-history.
At this stage there seems to be usually, if not always, cireu-
lation of the blood in the bases of the legs, but there is still none
in the antenne or in the lamelie. J am not certain as to when
circulation begins in these latter organs, but it appears to be
established about the 6th or 7th stage, if not earlier. One inter-
esting point concerning it is that it is apparently intermittent, as
there are times when it is impossible to see any corpuscles moving,
while at other times the flow is distinct and rapid. The circulation
in these organs seems to be confined to the region round the main
tracheal trunks, there being in each lamella a blood-space above
and below these trachee. The direction of the flow in these blood-
spaces seems to be reversible, as at times the blood may be seen
flowing outwards above the trachea and inwards below, while at
other times the opposite is the case. The circulation in the
middle lamella also is sometimes the opposite of what it 1s in the
lateral ones. At times the corpuscles oscillate backwards and
forwards on both sides of the tracheal trunks. I shall have more
to say concerning the lamelle when I come to discuss the moults.
In one individual examined at this stage there were three
Malphigian tubules.
Stage 4.
At the third moult no increase takes place in the number of
1909. | LIFE-HISTORY OF THE AGRIONID DRAGONFLY. 265
segments of the antenne. In the labium the labial palpi now
each bear 3 hairs, the new ones having been added below the
second. A second pair of hairs has also appeared on the body of
the labium between the other two hairs. These new hairs may
be as large as the first pair or they may be smaller, only attaining
full size at the next moult. In later stages the new hairs always
appear aS minute spines and are often only half their full size
after the following moult, although they are more usually full
size by that time.
A marked increase in the extent of the circulation is now
visible. ‘The corpuscles can be seen passing up the inner side of
the first, second, and part of the third antennal segments. About
halfway up the latter segment they cross over to the outside of
the antenna and return to the head. Circulation is also distinctly
visible in the bases of the legs as far as the trochanters, passing
out from the body anteriorly and back to the body along the
posterior side and beneath the tracheal tube.
In the femora an irregular movement of blood-corpuscles is
recognizable, but very few seem to be present and these are con-
stantly being temporarily held up among the muscle-fibres.
Usually, in this stage, small ridges appear on either side of the
second and third thoracic segments, this being the first trace of
wing-development. The appearance of these ridges is not, how-
ever, always in stage 4. In a few instances I have noticed them
in the previous stage, while in many cases they do not appear
until the fifth stage, but either stage 4 or 5 seems to be the
normal one for their appearance. ‘The development of the wings,
like that of the lamellz, seems more closely connected with body-
growth than with the moults. ‘This statement, however, requires
some qualification, for, although it is not possible to describe any
definite condition of wing-development as being connected with
any particular moult, there seem to be one or two very definite
stages correlated with the approach of the final ecdysis. This
will, however, be referred to later.
Stage 5.
A further increase in the number of antennal segments takes
place at the fourth moult, the third segment from the base again
dividing into two equal parts. This 6-segmented condition of the
antennz continues through the next four stages, in fact, until the
commencement of the final nymphal stage is reached, when a
change takes place.
The condition of the labium remains the same as in the previous
stage.
Although no regular record has been kept of the rate of
increase in the number of Malpighian tubercles, one nymph
examined in the fifth stage had either 5 or 6 of these. The rate
of increase in the number of these must become more rapid later
Proc. Zoou, Soc.—1909, No. XVIII. 18
266 MR. F. BALFOUR-BROWNE ON THE [ Mar. 2,
on, as Schindler (1878) describes 50 or 60 as being present in the
imago of Agrion puella.
Stages 6, 7, 8, and 9.
The labium in stage 6 usually has 4 hairs on each palp and
three pairs on the labium. Each new pair of hairs as it appears
on the labium arises between the original hairs, so that gradually
the series takes the form of a crescent with the ends curving
forwards stretched across the labium. In stage 7 the condition of
the labium is usually the same as in stage 6.
In stage 8 the labium may be just as in the previous stage, but
often, perhaps more usually, either a pair of new hairs has appeared
in the crescent or one hair only has appeared, so that there are
4 hairs on one side and 3 on the other. I noticed this irregularity
in a number of cases, and although such uneven developments
occur in other stages they do not seem to be so common as in this
one. Occasionally there are 5 hairs on each of the palpi. In
stage 9 there seem to be, with scarcely any exception, 5 hairs on
each palp and 4 pairs on the labium.
Stage 10.
This may be the final stage in the life of the nymph. The
antenne, if this is the final stage, will be seen to possess 7 segments,
the additional one having been produced by the division of the
6th immediately above the small spur already referred to. Iam
not certain that this division takes place in all the Agrionide,
but the statement holds good for Agrion pulchellum and puella,.
Pyrrhosoma nymphula, and Ischnura elegans. I think that the
nymph of Lrythromma naias has antenne of only 6 segments in
its final stage, but I can only find notes as to one specimen of this
species. Similarly, with regard to Hnallagma cyathigerum, the
only specimen of which I have kept a record had 6-segmented
antenne in its final stage. When this is the final stage of the
nymph, the labial palpi usually possess 6 hairs each, while the
body of the labium has a crescent of 10 hairs more or less evenly
spaced and all pointing forwards. In other cases, however, the
labium may remain as in stage 9 or there may be an intermediate
condition, 5 hairs on each palp and 5 pairs on the labium itself.
The blood-circulation in nymphs in the later stages is, in the
legs, apparently the same as described for stage 4. Bowerbank
(1833, p. 243) describes the blood as circulating in the leg of
Agrion “ even to the very extremity of the tarsus, where, as in
the antenne, the particles of the blood are seen to descend on the
one side of the leg and turning the extreme point to return up
the contrary side to the one by which they came down.” Even
in the full-grown nymph I cannot detect a regular circulation
beyond the trochanters, where the corpuscles can be distinctly
seen passing from the anterior to the posterior side and returning
1909. ] LIFE-HISTORY OF THE AGRIONID DRAGONFLY. 267
to the body. This circulation is rapid—as rapid as in the
abdominal segments,—but that in the tibie and tarsi is extremely
slow and irregular, the few visible corpuscles being constantly
held up for quite long periods.
In the antenne the circulation extends now to the apex of the
5th segment. A few corpuscles can be seen travelling up the
inner side very irregularly, there being constant delays and short
rushes. At the apex of the 5th segment they quickly pass over
to the outer side and travel back to the head in the same easy
stages.
Bowerbank (/. c.) mentions that the nymph he examined had
antenne of 6 segments and that the circulation only extended
to the extremity of the 4th segment—a condition probably
reached about the 6th or 7th stage, but I have not paid sufficient
attention to the circulation to be more precise.
Stages 11, 12, de.
If the nymph is not going to hatch out at the end of the 10th
stage, the antenne do not change from the 6-segmented condition.
It seems to be a rule in Agrion and /schnura that the 7-segmented
condition only exists in the final stage, while in Pyrrhosoma the
only two individuals which I reared through the last stages
showed the 7-segmented condition in the penultimate stage.
The labium may have either 6 or 7 hairs on each palp and 5 or
6 pairs on the body, while occasionally no further development
takes place after the 10th moult. If there are 7 hairs on each
palp they are very crowded and the 7th appears at the extreme
base of the palp.
The majority of my nymphs hatched out into imagines at the
end of the 10th, 11th, or 12th moults. Two Agrions passed
through 13 stages, while the two Pyrrhosomas passed through
11 and 12 respectively. A single Hnallagma passed through
14 stages as did Hrythromma, while an Ischnura imago emerged
at the 12th moult. I shall later on discuss the question of the
variation in the number of moults in nymphs of the same species.
Tue Movtts.
In the case of the Agrion nymphs which hatched from the
eggs at the end of August, the first moult took place in from 8 to
12 days. Their average length, after hatching, was 1:3 mm.,
while after the first moult it was 1-6 mm. The second moult
was a less regular event than the first, some nymphs undergoing
it 14 days later, while others took more than three weeks to
complete their second stage. The third moult again took place
from a fortnight to five weeks after the second, while the fourth
was even more irregular than the third.
At the beginning of November the nymphs become very
18%
268 MR. F. BALFOUR-BROWNE ON THE [ Mar. 2,
sluggish, showing but little interest in the Paramecia with which
they were abundantly supplied.
In the beginning of December, seeing that matters were at a
standstill, I placed 18 of the tumblers ina Hearson incubator and
maintained a fairly constant temperature of 19° C. (66°2° F.),
which isa very high summer temperature. The outer door of the
incubator was removed, so as to admit light, but, from the position
in which the incubator stood, the occupants had little more than
twilight even on the brightest days.
The effect of the warmth was soon obvious, as all the nymphs
in the incubator showed great activity and readiness to feed.
One nymph moulted 8 days after being placed in the incubator,
and others followed fairly soon. By the end of February several
of the nymphs had completed their 8th stage, while one was
through its 9th and another through its 10th. During the same
time the nymphs, some fifty and more, in the ordinary tempe-
rature of the laboratory did not moult at all, with the exception
of three specimens, two of which moulted at the beginning of
December and the other at the end of February. The average
length of the nymphs in the incubator was 11:0 mm., while that
of those outside was 2°7 mm. !
A large part of the experiment had now to be sacrificed, as I
was leaving Norfolk for Ireland, so the incubator was gradually
cooled off and the former occupants became less active and much
less hungry. On April 20th I started a similar incubator in
Larne, and this was kept going until July 10th, but only one or
two of the nymphs which had been kept warm during the winter
were placed in it, a number of the less advanced ones being
selected in order to further test the effects of temperature. The
new batch of nymphs quickly showed great activity and keen
appetites and began to grow rapidly, moulting at short intervals.
If further evidence were needed of the effect of temperature in
the causing of moults, it is to be found in the cases of three
nymphs, one in the 5th and two in the 6th stage, which had
stagnated for 92, 135, and 104 days respectively through the
winter. They were placed in the incubator for 21 days in
February. Each moulted 7 days after being warmed, and each
moulted a second time in the next 10 days. After being removed
from the incubator one moulted three weeks later, while the
other two were five weeks without changing.
This effect of temperature upon the duration of the stages
makes it impossible to find a fair average duration for any stage
except by using only those nymphs which have been incubated,
and only those stages passed through while in the incubator.
Unfortunately, this was not foreseen, so that the material at hand
is very meagre. I have, however, prepared a table showing for
19 nymphs the number of days occupied by each stage, indi-
cating the days passed in the incubator by using a thicker type
(v. Table 1).
From Table 1, reading down the columns and only using the
figures for stages begun and completed in the incubator, we can
Taste 1.—Showing the number of days oceupied by each stage of the nymph.
fen)
a The figures in thick type indicate that those days were spent in the incubator.
| | | Total |
Ret SrAGE! sonst Te IDG, | UNG IN We Wi 4} 0, |) wait, IX. XG XJ. | XID. | XIII.) XIV. | number
isa | of days. |
& | | |
5 | FE. | : |
) Nymph. Sex. |
BS A, pulchellum. Nala 12 17 ee 25 150 16+15 Pepa || 5 21 | 18+19 86 * 211 624,
=) » (2. yp wa) ilies 60* +19 20 8 | 14 | 20474) 41 296
a is +3, @..| 12 | 20 | 21 | 46+10 9 10 11 | 16 | +2 69 277
= F WA, Bacall 12 17 14 | 57+12 9 12 10 14 14 10+48 x 229
S) : 5B @..| 12 | 24 | 82° | Leer? 9 31 145 23 2 epep eee 50) || 634
f= - qh i. 7 Ni/ 14) 25 92+7 9 6+21 | 31+15 Wel 24 25+29 | ... 339
o i Wo — Reo I 24. 55 | 24+15 4 10 en | i} | 9 22 6+81 | 37 om. 291
= e +3. @ ?+15| 18 9 14 he Bits lt 4+8 2 26 By | Daly)
A ion Oe | Seca ape esestal 17 14 21 18 | 40 49 227
a 2 SHPO, aac tb) Tae a oe all 28 135 +7 | 6 | 8+388 | 16+11 36 |
S » SPU ver | 14 | 23 26 | 104+7|) 8 1437 | te+7 | 92 | 5e-12| |
© 3 SR a noel | 18 7 35 133 | 388 |16+18; 14 49+8 | | |
Be " IEE hes FP 14 28 |60+10| 9 10 14 17 =| 21+37 | 69 |
is 3 i, Oy = 13 14 a) | il 2410 | 17 24 «| 804+37 | 71
& IT, elegans. NS 1 cenlf aes ce | ol 21 34 26+8 | ae al) 9 22 8+?
= |r. naias. UGS nal eave Ghat | oaks ine 28 60+19 | 20 18 8 13 8+2 | 51 37 56
Ht | Bn. cyathigerum. $17. ? ... leer 7 og TBEY | Gael ale 12 44 |2+63| 216 | 30 |
H |P.nymphula. 18. ?... | ae) 23 32 | 28+14; 18 | 18 13 21. 7474 |
5 ‘¢ ig), F 13 ae) |) essa) RY I) ie) 16 | 25+18| 983
ae * A moult was by mistake not recorded; in No. 2 it occurred in the incubator.
en + Only in incubator from April 20th onwards; not during the winter.
= + Only in incubator during the winter months—December to February.
ei
§ Incubated for 21 days in February, otherwise no winter warmth.
270 MR. F, BALFOUR-BROWNE ON THE [Mar. 2,
find the average number of days occupied by each stage, and the
results, so far as it is possible to get any, are as follows :—
TABLE 2.
arya Number Max. Min.
Stage 5 of number of | number of
number of days. nymphs. days. days.
Vv. 9°5 6 ia t
VI. 15 8 31 9
VII. 12°0 14 22 6
VIII. 133 13 23 5
IX. 14-1 12 21 8
Xe 19°2 10 AT + 9)
Nal 29°44 5 52+ | 10 ;
The figures indicate that the stages tend to lengthen from the
earlier to the later ones. There is, however, a source of error in
the method used, and that is that while some nymphs are full-
grown at the end of the tenth stage, others go through 11, 12, 13,
or 14 stages. Now one point which is quite clear without any
statistics is that the final stage is always a fairly long one, and
this fact will of course affect the above figures. I have therefore
checked these results by using Table 1 in another way. I have
taken the average duration of the last stage of each nymph, then
of the penultimate stage, then of the one before that, and so on as
far back as my figures will permit, again of course only including
stages begun and completed in the incubator, and the results are
as follows :—
TABLE 3.
| | Penultimate Last
: Stage. Stage.
Successive i A ee 2
stages ; 4—8 5—9 6—10 7—11 8—12 | 9—13 10—14.
Number of ; |
nymphs } 4, 11 14, 15 iL 6 3
Average no. 12°3 10°7 13°4 147 | 168 19°5 456+
of days |
Max. & min. 2 | / / i 4 FS 8
no. of days $ ly, 10, 22, 8 |) Sil, &, | 44. 6. | 24. 9. 24, 14. 62+ 36.
Again the figures show that the stages tend to lengthen out as
the nymph grows bigger.
The first imago to appear emerged 33 weeks (230 days) after
hatching from the egg and after passing through 10 stages. This
was one of the individuals which spent the winter in the incubator,
and I expected to find that all the nymphs which had been kept
warm during winter and had fed and moulted regularly would
complete their life in the same number of stages. However, out
of ten such nymphs only three got through in 10 stages, one in 11,
five in 12,and one in 14. The three nymphs which completed in
10 stages were placed in the incubator in their 4th stage, aS was
1909.4) LIFE-HISTORY OF THE AGRIONID DRAGONFLY. 271
also one which passed through 12 stages, while all the other
nymphs were in their 6th, 7th, or 8th stages at the beginning of
December. Now all these nymphs were in the incubator at the
same time and for 81 days, and during that time they moulted
a number of times, varying from 3 to 7. If we consider the
number of times that each of those nymphs moulted in relation
to the total number of moults it passed through, we find that
those introduced into the incubator in stage 4 passed through half
(5 out of 10 and 7 out of 12) their total number of moults in the
81 days, while those introduced at later stages passed through less
than half their total number—a fact which supports what I have
already shown by two other methods, namely, that the stages tend
to lengthen out as the nymphs advance.
Whether or not this lengthening out of successive stages is a
phenomenon common to the larvee of all insects has still to be
shown, and I can find no reference to the subject, but it is
interesting to note that Newport (1839) mentions that the rate
of the pulse in the larve of Sphinw ligustri gets slower in
successive stages—a fact which might well be correlated with the
lengthening of the stages.
Leaving out for the present the question as to whether the
winter warmth has curtailed the number of the stages, a point on
which I have very little evidence, it is easy to show that the
favourable conditions greatly curtailed the duration of the life of
the nymph.
All the imagines from the winter-warmed nymphs appeared
between April and July of the year following that in which the
eges were laid—that is, within 10 months. On the other hand,
the imagines from the nymphs which were in ordinary laboratory
temperatures during the winter, with one exception, appeared a
year later than the others, more than 20 months after the hatch-
ing of the eggs, and in spite of the incubation from April to July.
In connection with these two extremes, the three cases of nymphs
which for 21 days in February enjoyed the warmth of the
incubator are most interesting. In all other respects they were
treated exactly like the second batch of nymphs—that is, they
endured ordinary laboratory temperatures until April 20th, and
then were in summer temperature,—yet they all three hatched out
with the first batch, one in June, one in July, and the third in
August. It is certainly interesting that three weeks’ warmth in
February, during which each nymph moulted twice, should have
made a difference of more than 10 months to those three nymphs ;
and since temperature has such a marked effect it may be possible
to account for the periodic swarms of dragonflies on the ground
that a specially mild winter in some region may bring ona batch
of nymphs, so that the imagines appear a season earlier than under
normal circumstances. In such a case the batches of 2 years
would appear together, and might be a prelude to a migration.
I cannot account for the single individual which, without any
winter warmth, emerged with the first batch of nymphs. It is
possible that it was in the incubator with the other three in
[ Mar. 2,
MR. F. BALFOUR-BROWNE ON THE
272
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1909. ] LIFE-HISTORY OF THE AGRIONID DRAGONFLY. 273
February, as it moulted once during that period, but so far as my
records go it was not !
Table 4 (p. 272) gives the dates of appearance of the imagines
and also the number of stages through which the nymphs
passed.
With regard to the question whether a high temperature in
winter tends to reduce the number of stages in the life of the
nymph, the material is too scanty to form any sound opinion.
For such a question only one species can be considered at a time,
as probably all species do not moult the same number of times
under similar conditions. Also the individuals of any one species
will not, under apparently similar conditions, all moult the same
number of times, so that the average of a fair number of
individuals would be necessar y to get any definite result.
With regard to Agrion pulchellum, the only thing that can be
said is that, with the single exception already referred to, the
only nymphs which completed in 10 stages were three w hich had
been incubated during the winter, while the only ones which
passed through the ie stages were two which had been kept cold
during the winter. On the other hand, three of the winter-
warmed nymphs passed through 12 stages, while two of the others
got through in 11; andlam inclined to think, after studying the
rate of growth, that the incubator did nothing towards reducing
the number of stages. I shall discuss this in relation to the
question of body-length.
Bopy-LENGTH.
The average length of a nymph of Agrion pulchellum in its
final stage is 18°0 mm., the smailest nymphs being about 14-0 mm.
and the largest about 22°0 mm. These figures were obtained by
measuring a number of nymphs with 7-segmented antenne
collected without any regard as to the number of stages they had
passed through, and they show a wide range of variation in size.
In the newly-hatched nymphs there was but little difference in
body-length, but at each succeeding moult the differences became
more marked, so that, for instance, after the eighth moult, two
-nymphs measured respectively 6°7 mm.and 14-0 mm. In noting
this fact I also noted that the smaller nymph passed through
13 stages, while the larger one completed in 10; and as the latter
had been incubated during the winter, while the former had not,
I at first thought that there would be evidence of reduction in
the number of moults in the winter-incubated individuals. To
test this I took the body-lengths of all the nymphs, so far as I
had the figures, at each stage, and compared the length of each
nymph in relation to the number of moults it ultimately went
through, and the results are rather interesting and are given in
Table 5(p. 274). It will be noticed that the tendency all through
is for the larger nymphs in any stage to be those which ultimately
complete in the smallest number of stages; and since this is
noticeable in the beginning of the second stage and becomes
more marked in succeeding stages, it seems that the earlier stages
[ Mar. 2,
MR. f. BALFOUR-BROWNE ON THE
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1909. ] LIFE-HISTORY OF THE AGRIONID DRAGONELY. 275
of the nymph decide its destiny as to the number of stages.
Incubation, then, had apparently no effect on the number of
stages which had been determined before incubation was resorted
to, but it seems probable that if the nymphs had been incubated
from the moment they hatched, the effect would have been
different. That there is not complete regularity in the Table is
perhaps not to be wondered at. A nymph may “ stagnate” for
a period owing to some upset of its metabolism, and that might
cause it to fall behind for a stage or even longer, in which case
the question arises whether a long delay would not possibly
necessitate an extra moult? I will consider this point further
when discussing the significance of the moult.
THE ANTENNA.
The segmentation of the antennz has already been described.
It is to be noted that the segment immediately above the scape,
i. e. the third from the base, is the one which divides three times
to produce new segments for the antenna, and that these divisions
are, with a very few exceptions, definitely connected with certain
stages in the life of the nymph. The actual formation of the
new segment is not, of course, an instantaneous process, but takes
place by slow degrees within the “shell” of the antenna, only
becoming revealed when that shell is cast off.
It is interesting to note that in one of the Ephemeride, where
development has been followed, Lord Avebury (1863 and 1865)
found that the third segment of the antenna, the one next to the
scape, is there also the centre of growth.
As the nymph increases in length the antenne also grow, but
these organs do not maintain a rate of growth proportionate to
that of the body. Whereas in the first stage the antenne are
23 per cent. of the body-length, the percentage gets steadily less
as the nymph grows, until, in the full-grown individual, the
antenne are less than 12 per cent. of the body-length.
The growth in length of the antenna is not due to the gradual
lengthening of each segment, but is chiefly due to growth of the
region immediately above the scape, This is easily shown by
taking the average length of the antenna at each successive
stage and finding what percentage of that length is taken up
by each segment. The results are shown in the accompanying
chart (p. 276), from which it will be seen that, whereas segment 3
gradually increases its percentage of the length of the antenna,
the scape and segments 4 and 5 only maintain a rate of growth
proportional to that of the antenna, while segment 6 grows much
too slowly to maintain its proportion to the total length.
Stages X., XI., XIT.,and XIII. may any of them be the last stage
in the life of the nymph, and in finding the average length for each
segment at each stage I had to exclude the “ 7-segmented ”
wa eee in each of these stages, except XITI., where there were
“¢§-seomented” nymphs. For comparison I also found the
meee for the “ 7-segmented” condition in stages X., XI.,
and XII., and they are given as X.a, XI. a, and xan ce: and it
will be seen that in each of them segment 3 is proportionately
276 MR. F. BALFOUR-BROWNE ON THE [ Mar. 2,
longer than in the ‘6-segmented” condition. The average
length of the “7-segmented ” antenne is also greater than that
Chart showing the growth of the segments of the antennex in Agrion pulchellwin Lind,
of the ‘‘6-segmented” antenne for the same stage, indicating
that the larger nymphs of the stage are those which are near
their final ecdysis.
THe LAMELL2.
Like the antenne, the lamellz are much longer in proportion
to the body-length in the newly-hatched nymph than in the full-
grown one. Whereas they are almost the same length as a
nymph of 1°3 mm., in a nymph of 17 mm. they are less than
half the length.
The pointed linear organs of the first-stage nymph gradually
widen out in the vertical plane and ultimately become leaf-like.
In both Agrion pulchellum and puella, and perhaps in all species
of this genus, they gradually lose their pointed appearance and
become distinctly rounded at the apex. Not only have I observed
this rounded character in nymphs reared from the ege, but I
have repeatedly hatched out the imagines of both these species,
and only those nymphs with rounded lamelle produced Agrions.
I mention this because Lucas (1900, pp. 281, 288) describes the
lamellee of A. puella as being always pointed, and says that those of
A. pulchellum are appar ently similar. He mentions, however, that
some authors have described them as having rounded apices. The
full-grown nymph with pointed lamelle, which is otherwise almost
(or quite 2) indisting wishable from the Agrions, is [schnura elegans.
1909. | LIPE-HISLORY OF THE AGRIONID DRAGONFLY. 277
The lamellee of Agrion—I have been unable to find a character
to distinguish the nymphs of A. pulchellam and puella from one
another—are indistinguishable at first from those of Jschnura,
unless possibly by certain small patches of pigment; but about
the 6th, 7th, or 8th stage, sooner or later, according to the size
of the nymph, they have become sufficiently wide and rounded at
the apex to distinguish them,
These organs, into which the lateral tracheal trunks of the
body pass and in which they break up into minute branches, so
that the whole lamella is filled with them, are said to have ¢
respiratory function. I have already referred to the ianeul
mittent blood-stream in these organs. Whether it is correct to
describe it as intermittent I am not sure, but I have several
times noticed that in a nymph taken from the tumbler and
placed under the microscope no circulation was to be detected
until after the lapse of some time, and it has occurred to ine that
the circulation ih the lamellse was noticeable only when the air
in the water was becoming exhausted.
These organs are not, ‘however, indispensable, as the nymph
can live quite well without them. In the final stage they are
often without them, having apparently cast them off. In the
earlier stages, however, if a lamella is removed it is replaced by
degrees. If two or three days previous to a moult a lamella is
removed, no replacement takes place until the second succeeding
moult. As soon as the nymph escapes from the old skin, a very
thin pointed sac, more or less circular in transverse section, is
visible in place of the lamella previously removed. This sac is
much smaller than the other Jamelle and contains at first no
trace of trachee. It takes two, three, or even four moults before
a new lamella is indistinguish: ble from the uninjured ones. The
process of renewal is not "sudden as it appears to be. The wound
caused by removing a lamella is apparently quickly closed and
the formation of the new lamella commences within the “ shell,”
I believe at the time when a new nymph-skin is forming within
the old one. Presumably ib forms in a crumpled condition like
the imaginal wings within the nymph wing-cases, and as soon as
the pressure of the outer shell is released, the new lamella expands
asa sac, becoming turgid owing to fluid enteri ing it from the body.
A leg i is replaced i in the same way as a lamella, taking two or
three stages to attain perfection, appearing immediately after the
moult as a thin-walled sac which gradually thickens and hardens.
I have paid very little attention to the replacement of lost
parts, but it seems to involve two processes: first, the formation
of a thin-walled sac in the form of or tending towards the form
of the organ to be renewed, and, secondly, a re-formation within
the sac of the muscular and tracheal systems,
WING DEVELOPMENT,
I have already mentioned that the first sign of the pleural
ridges appears after the nymph has completed its third or fourth
278 MR. F. BALFOUR-BROWNE ON THE [ Mar. 2,
moult. This statement is true of Agrion, Ischnura, and Pyrrho-
soma, and may be true of Hnallagma and Hrythromma, but I
imagine that in these cases wing-development begins rather later.
These ridges lie diagonally across the second and third thoracic
segments on either side of the tergum, and are the result of the
upward growth of the pleura of the meso- and metathorax.
The rate of development from the first differs somewhat in
different individuals. In the stage succeeding that in which the
ridges first appear there may be a small bud on the inner side of
each ridge, representing the commencement of the “wing,” or
this may not appear until a stage later.
As the buds grow larger the ridges also develop, those of the
mesothorax gradually spreading anteriorly over the tergum until
they meet in the middle line. Posteriorly they do not approach
one another, so that a triangular patch of the tergum is left
exposed between them. The metathoracic ridges joi on to the
posterior edges of the mesothoracic ones, and taper away
posteriorly towards the sides of their segment, the posterior
margin of which they slightly overlap. Thus a large part of the
tergite of the metathorax is left exposed. The “ wings” develop
in such a way that they lie more or less flat on the sides of the
dorsum in the wedge-shaped depression between the ridges, and
extend backwards over the anterior abdominal segments. Towards
the end of the final stage, however, when the thorax of the
imago is forming within that of the nymph, the “wings” are
somewhat raised and tilted.
At each moult an advance in development is visible, but it
differs very considerably in the earlier stages in different indi-
viduals.’ For instance, a condition when the anterior ‘“ wings”
just overlap the posterior edge of the metathorax, and the
posterior wings just cover the first abdominal segment, might
represent the 8th, 9th, or 10th stages, apparently depending on
the general body-growth of the nymph and the number of stages
through which the nymph is going to pass. There is a distinct
connection between the later moults and the proportion of
increase in the length of the wings. For instance, if the anterior
wings reach in one stage almost. to the posterior margin of the
metathorax, their increase at the next moult will be about
33 per cent. of their length. At the next moult the increase
will be about 40 per cent., while at the next, whether the 10th,
11th, or 12th, &e., the increase will be about 50 percent. This
is a large increase, and is, in every case I have observed, evidence
that the nymph has reached its final stage. The “ wings” now
cover the first three abdominal segments, and often extend half-
way over the fourth, and any nymph found with such long wing-
cases is in its last and generally its longest stage.
The last nymphal stage is characterized by these long “ wings”
and by the 7-segmented antenne, and the one or two stages
previous to that are fairly well-marked by the rapid growth of
the wings. The first four stages in the life-history are also easily
recognized by the state of the antenne and of the mask. In my
1909. } LIFE-HIStORY OF THE AGRIONID DRAGONFLY. 279
experience it was, with only one or two exceptions, after the
antenna had become six-segmented that the nymphs showed such
variation in their rate of progress even when under similar
conditions.
RESPIRATION.
The caudal lamellae have been described as respiratory organs,
and have been called “ caudal gills,” but Dewitz (1890) pointed
out that the nymph can live without them, and suggested that
possibly a rectal respiration, similar to that of the Calopterigide
and the Anisopterids, existed in the Agrionide also. In those
forms, however, where rectal respiration is dominant, there is an
elaborate arrangement of trachee in the walls of the rectum
which is completely absent in the Agrionide.
Dewitz has shown that a stream of water does pass in and out
of the rectum in the Agrionide, but from my observation it is a
very weak one, nor is there any special apparatus surrounding
the anus in this group to prevent ingress of foreign particles
such as is found in the Anisopterids. If rectal respiration exists
at all, it seems to me that it must be very slight and of but little
importance, as I could not observe any increased number of
contractions of the rectum in specimens of Agrion which had
been deprived of their lamellee.
In the absence of the lamelle, I thizk the whole of the
respiration must be carried on through the skin,
CONCLUSION.
The occurrence of a pronymph stage in the Odonata * is not
unique. It has been observed by Pagenstecher (1864, p. 7) in
Mantis religiosa, and occurs also in other Orthoptera [see Kiinckel
(1890 (1) p. xxxvii, and 1890 (2) p. 657)], and in several other
orders (Packard, 1898, pp. 583-585), and is no doubt more
common than has at present been recorded.
The stage is apparently for the purpose of freeing a tightly-
packed larva from the egg, the amnion, instead of breaking and
remaining within the shell, continuing intact until after the
larva is clear, and the amnion forms the pronymph skin.
In the Odonata the larva casts the pronymph skin in exactly the
same way as it casts its skin in subsequent moults, and it appears
that, in the other forms in which this stage has been observed,
the amnion is got rid of in much the same way, except that, as a
rule, a special ampulla exists in the thorax of the larva which, by
swelling up, bursts the amnion in that region (Packard, /. ¢.
p. 984).
Although the total number of moults may vary somewhat,
there is an unmistakable connection between certain stages in
the development of the nymph and certain moults. What I have
said as to the growth of the antenne, the appearance of the labial
hairs, and the development of the wings shows this, especially in
* Exactly the same phenomenon occurs in Sympetrum striolatum, one of the
Anisopterid dragonflies, where the pronymph stage continues for about 7 minutes.
280 MR. F. BALFOUR-BROWNE ON THE [ Mar. 2,
the earlier stages, and it is possible that a large number of
nymphs kept under uniform conditions might have caused the
correspondence to have continued through every stage.
The correspondence between certain stages in development
and certain moults has been observed in the larve of other
insects. For instance, Acrydians moult five times, the wings
appearing after the second moult and i increasing quite definitely
at each succeeding moult, the merease in the “last stage being
very great. “ Besides the acquisition of wings there are slight
differences at each moult both im structure and colour, besides
size, so that we may always recognise the comparative age and
the particular stage of growth of any individual” (Packard, bt:
9. 595). Tord Av bury (J. ¢. p. 62) noticed that the nymph of
Chloéon “ oradually attains its perfect condition through a series
of more than 20 moultings, each accompanied by a slight change
of form,” and there are numerous other cases cited in works on
entomology, but, by many authors, no notice seems to have been
taken of these facts in discussing the question of the significance
of the moult.
On this latter point various suggestions have been made, of
which the simplest, and perhaps the oldest, is that the moults are
necessary to enable the larva to grow. It is obvious that where
a larva is encased in a hard intesument, one of two processes is
necessary in order that the contained organism may Increase in
size. Hither the integument must be capable of expanding with
the organism, or it must be cast off from time to time.
When the nymph moults it expands greatly in size as it creeps
out of its old skin, and, a few minutes after escaping, is much
larger than the agin from which it emerged. The nymph has
therefore been in a state of compression within its skin previous to
the moult. But growth does not only take place at the moment
of the moult—for instance, a nymph measuring 1] mm. at the
commencement of its final stage was 15°2 mim. at the end of it,
so that the nymph-skin is undoubtedly capable of expansion.
On the other hand, Dr. Sharp (1895, p. 163) considers that
growth is only of secondary importance in accounting for the
moult, urging as one reason that ‘“‘many insects make their first
ecdysis almost immediately after they leave the egg”; and he
quotes the case of the blowfly larva, which, “ according to Lowne,
moults within two hours of its emergence from the egg.” The
case of the dragonfly pronymph is even more apt in this connection:
Hisig (1887, p. 781) suggested that the moult was for the
purpose of getting rid of w aste nitrogenous matter, and Dr. Sharp
(U. ¢. p. 163) adds that it may be of importance in separating the
carbonaceous products from the blood. Bataillon (1892, p. 61;
1893, p. 18) has stated that at the period of the moult the larva
is inastate of asphyxiation ; and Dr. Sharp (/.c. p. 131), referring
to this observation, remarked that if it is a fact, ‘‘ the secretion
of the new coat of chitin may figure as an act of excretion of
considerable importance.”
The method of passing to the integument certain waste
1909. | LIFE-HISTORY OF THE AGRIONID DRAGONFLY. 281
products of the metabolism is almost universal throughout the
animal kingdom, aud there seems no reason why it should be
regarded as, in insects, a special process of excretion. Whatever
may have been the primary cause, the process has apparently been
seized upon by Natural Selection, which has eliminated the less
protected individuals, and produced a tough protective exo-
skeleton.
Bataillon’s observations do not seem to me as important as
Dr. Sharp regards them in demonstrating that the moult is a
special act of excretion. When it is remembered that the larva,
immediately after a monlt, is soft and helpless, unable to feed,
and an easy prey to almost any hungry animal, it is possible to
believe that those individuals which did not quickly produce a
new protective armour were eliminated in the struggle for
existence. In order to quickly produce a new hard skin, a larva
would necessarily retain a large amount of waste products which
would otherwise pass out of the system, and this might account
for an asphyxiated condition immediately prior to the moult.
Pantel (1898) considered the moult as necessary to account
for the formation of ectodermal organs, and points out that, at
each of the three stages of the larva of Thrixion, new characters
appear; while Packard apparently held much the same view,
as he says (/. c. p. 615), “the swollen bodies of the gravid female
of Gastrophysa, Meloé, or of Termites and of the honey-ant show
that the skin can stretch to a great extent, but in the meta-
morphoses of Crustacea and of Insects, the young of which are
more or less worm-like or generalised in form, with fewer segments
and appendages, or with appendages adapted for quite different
uses from those of mature life, the necessity for a change of skin
is seen to be necessary for mechanical reasons. Hence Crustacea
and Insects moult most frequently early in life, when the changes
of form are most thoroughgoing and radical, while simple growth
and increase in size are most rapid at the end of the larval life,
as seen in both shrimps and crabs, and in insects.”
Now it seems to me that the fact that the larva is in a state of
compression within the old shell at the time of the moult,
suggests that the moult is necessary for growth, while the fact
that certain stages of development are correlated with certain
moults suggests that the moult is necessary for development, and
Tam inclined to think that a moult may be either a “‘ growth-
moult” or a ‘‘ developmental moult,” or a combination of the two.
Growth and development are, to some extent at least, inde-
pendent phenomena, since we find variations in size in equally
mature individuals, and apparently either growth or levelopment
may be retarded without the other being affected, as any rate to
the same extent.
From the fact that certain stages in the life-histery of a larva
are correlated with certain advances in development, growth
cannot possibly control the moults producing those stages, and
they must be primarily ‘developmental moults.” Further,
Proc. Zoou. Soc.—1909, No. XIX, 19
282 MR. F. BALFOUR-BROWNE ON THE [Mar. 2
assuming that there are ectodermal developments to take place,
it seems to me that so long as those developments do not take
place in stages of excessive length, development will always be
the contr olling factor.
From the time of the moult the new ectoderm of the nymph 1s
steadily hardening (probably oxidizing), and gradually losing its
elasticity. As soon as the skin has ceased to expand with the
growth of the nymph, the pressure on the body of the nymph
reacts so that a new layer of chitin is laid down within, and
separate from, the old layer. Once this new layer has completely
isolated the old one the latter becomes brittle, and gives way
along a line of weakness in the median line of the thorax when
the nymph exerts itself and escapes. Such would be the explana-
tion of a ‘‘growth-moult” pure and simple. If, however.
previous to this condition being reached a developmental stage
had been arrived at, a moult will take place which will serve
both purposes, but will have been caused by developmental
requirements.
In the case of one of my nymphs two moults took place within
4 days of each other. The second moult could not have been
required for growth nor, on the excretion theory, for excretory
pu pOSes, and I imagine that the first moult was a “ growth-
moult ” and the second a “developmental” one, especially as the
first moult was the first to take place after the nymph had been
put into the incubator, and 17 days after the previous one.
In this connection the effects of temperature on the number of
moults are important. Packard (/. c¢. p. 616) refers to the
experiments of Weniger and others on the larve of certain
Lepidoptera, the results of which show that under warm moist
conditions the number of moults is reduced from 5 to 4; and
referring to certain other experiments, he says (p. 618) : 7 ‘As to
the cause of the great number of moults in the arctians and in
the beetles exper mented with by Riley, it would seem that cold
and lack of food during hibernation were the agents in arctians,
and starvation or the lack of food in the case of “the beetles, such
cause preventing growth, though the hypodermis-cells retained
their activity.” W.H. Edwards states that larvee of Lepidoptera
which hibernate moult more often than those which have only a
summer existence, and larvee with a wide geographical distribu-
tion moult more often in cold than in warm regions. He also
says “there seems to be a necessity with hibernators of getting
rid of the rigid skin in which the larva has passed the winter ;
that is, if hibernation has taken place during the middle stages,
as it does in Apatura and Limenitis. In these cases, very little
food is taken between the moult which precedes hibernation and
the one which follows it, and the larva, while in lethargy, is
actually smaller than before the next previous moult. The skin
shrinks, and has to be cast off before the awakened larva can
grow” (quoted from Packard, /. ¢. p. 615).
1909.) LIFE-HISTORY OF THE AGRIONID DRAGONFLY. 283
Such a moult as that described by Edwards is apparently a
“ srowth-moult ” pure and simple. Unfortunately, however, it
is not always so easy to determine the nature of a moult
Development involves a certain amount of local growth which
presumably proceeds steadily between the moults. Ifa “ growth-
moult ” is interpolated, the ectodermal structures will noes remain
as they were before the “ growth-moult,’ but will show some
advance as the result of that moult, so that, if in the ideal
individual there are a definite number of developmental stages
each showing a definite advance, the interpolated ‘“ growth-
moult” upsets this state of things. But the developmental
stages must get right again by the end of the larval period,
so that there must be some power of adjustment; in other
words, the interpolation of a single “ growth-moult” will not
necessarily add merely one moult to the life-history, but may
necessitate one or more additional developmental moults for
adjustment. The more normal moults in the life-history, the
more probable it will be that the developmental stages will be
complicated by stages for adjustment. The lengthening of the
later stages of larval life, which Packard (J. c. p. 615) regards as
chiefly growth-stages, seems to me to be a lengthening of the
developmental stages owing to the more radical changes which
they involve. The last stage of the nymph, for instance, in which
the most marked internal changes take place is always a long one
and corresponds with the pupal stage of the Holometabola.
The fact that certain moults have nothing whatever to do with
growth (e. g., of the pronymph of the dragonfly; of the sub-
imago of Ephemerida ; moult to pupa and imago, &e.) seems fatal
to a theory that growth is the cause of the moult, while the fact
that certain moults have apparently nothing to do with develop-
ment (e. g., the moult of larve after hibernation) seems fatal to
a theory which makes development the cause of the moult.
Since, however, both causes seem to be at work, it is possible
that both theories, combined and readjusted, may give us a true
explanation of the significance of the moult.
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des Schmetterlings von Bombya mori.” Zool. Anz. x.
pp. 562-3.
Suarp, D. 1895—Insects. Cambridge Natural History, v.
London: Macmillan & Co.
Wueeter, W. M. 1891-3.—‘‘ Concerning the Blood-tissue of the
Insecta.” Psyche, vi. pp. 216-220.
Wueeter, W. M. 1891-3.—‘The Primitive Number of Mal-
pighian Vessels in Insects.” Psyche, vi.
1909. | ON GROWTH-STAGES IN BRITISH SPECIES OF CORALS. 285
EXPLANATION OF THE PLATES.
PratEe XXXII.
Development of Agrionid Dragonflies.
Fig. 1. The egg of Ischnwra elegans Lind., some hours before hatching.
Fig. 2. Part of the same egg, at a later stage, showing the shell-rupture and the
vesicle formed, and the head just commencing to move forward.
Fig. 3. Part of the egg, at a still later stage, showing the head X almost filling the
vesicle. a is the point at which the membrane will rupture for the escape
of the pronymph.
Fig. 4. The pronymph immediately after its escape from the ege.
5. Prothorax, dorsal view, of the imago of Agrion puella L., 9.
Fig. 6. Prothorax, dorsal view, of the imago of Agrion pulchellum Lind., °.
Prate XXXIV.
Development of Agrionid Dragonflies.
Figs. 1, 2, 3 show the last stages of wing-development in the nymph of Ischnura
elegans Iand., in this case stages 10, 11, and 12. The drawings are from
the moulted skin.
Fig. 4. The “mask” of Agrion pulchellwm Lind., in the first stage, upper side.
Fig. 5. One of the labial palpi of the same “mask” X about 400.
Fic. 6. The “ mask” in the third stage.
Fig. 7. The “mask” in the seventh stage, showing the palpi almost closed down on
the anterior border of the labium. The stiff hairs are so situated that when
the nymph is feeding they assist in holding the food, the mouth being
situated jurt bove b.
4. Growth-Stages in the British Species of the Coral Genus
Parasmilia. By W. D. Lane, M.A., F.Z.S8., British
Museum (Nat. Hist.).
[ Received February 16, 1909. |
(Text-figs. 38-46.)
I. Carruruers’ “ PHASES’ IN CARBONIFEROUS CoRALS.
In the early numbers of last year’s ‘ Geological Magazine’ an
article by R. G. Carruthers appeared on some simple Carboni-
ferous Corals*. In it stress is laid on the necessity of cutting
serial sections of corals and for the determination of species
according to their growth-stages. It is shown that Carboniferous
corals previously described as various species of Zaphrentis are
really different forms of Caninia cornucopie Michelin, and that
during development five “ phases” can be recognised, occurring in
the following order: (1) that corresponding with Zaphrentis ver-
micularis de Koninck; (2) with Lophophyllum dumonti Edwards
& Haime; (3) with Zaphrentis nystiania de Koninck; (4) with
Zaphrentis edwardsiana de Koninck ; (5) with Caninia cornu-bovis
Michelin.
It is important that the term ‘“ phase” is used to designate
these forms and not the implicatory “ stage,’ and the reason is |
* R. G. Carruthers, 1908, Geol. Mag. pp. 20, 63, & 158.
286 MR. W. D, LANG ON GROWTH-STAGES [ Mar. 2,
given :—“Save for the appearance of the dissepimental ring (a
feature confined to the mature part of the corallum) none of the
phases above described is truly ontogenetic, for, as will now be
explained, they are intimately connected with the habits of growth
of the corallum.” This is only an expression of the common difti-
culty in the determination of species of any group, especially
among fossils, the environment of which when alive is so
uncertainly understood—that of knowing whether a given cha-
racter 1s directly the result of heredity or caused by environment,
These “habits of growth” are shown to be the external shapes
of the corallum at a given size, two shapes being mentioned,
Trochoid and Cylindrical. And it is obvious from the examination
of simple corals that a Trochoid or a Cylindrical shape-stage 1s
always present, and if both, the former always precedes the
latter. For simple corals, then, two definite shape-stages may be
postulated ; nor can the environment directly reverse the order
in which they oceur. Other growth-stages in the shape of the
corallum may be found in simple corals; for instance, an Inverse-
trochoid shape-stage occurs in some Montlivaltia following the
Cylindrical shape-stage, and a Discoid shape-stage in other M/ontli-
valtia as an episode in the Trochoid ; but these need not here be
considered.
So far as Caninia cornucopie is concerned, it appears that the
environment determines the size reached by the corallum before
it passes from the Trochoid to the Cylindrical shape-stage. If this
were all, Carruthers’ “ phases” in C. cornucopie might be adopted
unhesitatingly as “truly ontogenetic” growth-stages, and the
determination of the species only complicated by the wide limits
of variation of the diameter, so that a comparatively large spe-
cimen might be found in an early growth-stage, and vice versa.
But it is stated that “it often happens that examples showing
the long septa of the dwmonti-phase are found to have the am-
plexoid septa,” characteristic of the next—the nystiana-stage—
“‘in their lower portions.” This appearence of an older stage,
earlier than, that is proximal to, a younger one, might seem to deny
that the “ phases” are growth-stages. The sequence in which the
“phases ” occur, however, is nowhere stated to be irregular, that is,
haphazard and different in different individuals; on the contrary,
except for the above modification, it is implied that the order of
phases is in regular sequence. An explanation of the apparent
irregularity is suggested by the behaviour of the coste in the
English species of Chalk Parasmilia, which alter in their orna-
mentation and general shape in a definite way when traced from
the proximal to the distal end of the corallum.
Before describing these it may be said that Carruthers mentions
other difficulties in the way of accepting his “ phases” as true onto-
genetic growth-forms. It appears that some internal characters
change in correspondence with the change in general shape, others
with change in the actual size of the corallum. Consequently, in
extreme cases of non-correspondence between the size and shape
1909. | IN BRITISH SPECIES OF CORALS. 287
of the corallum, some internal characters at a given time will not
be at the stage of development corresponding with the expected
synchronous stage of the other internal characters. How far this
is the case and how far it may be explained in the same way as
the irregularity in the sequence of the costal ornamentation in
Parasmilia, further work will show.
IT. GrowtH-Staces 1n CHALK PARASMILIA.
The English species of Chalk Parasmilia are attached by their
proximal ends to foreign objects. That this is an environmental
rather than an hereditary character seems probable from the
freedom and varying degrees of fixation in Montlivaltia, pre-
sumably of one species and from the same horizon and locality ;
for example, in J. rugosa Duncan *, from the armatus-zone of the
lias of Honeybourne, E. of Evesham, Worcestershire. It is con-
ceivable that on a muddy sea-bottom a simple coral would find
support in the mud around it, sufficient to hold it upright, while on
a hard bottom it would have need to be fixed. The environment
of the Chalk sea evidently required the Parasmilia to be attached
to some hard object. Owing to their fixation, the proximal end
spreads out to form a surface of attachment. So there is in
Parasmilia, first a Peduncle shape-stage, next a Trochoid, and
ultimately, in most, a Cylindrical shape-stage.
The English species of Parasmilia which have been up till now
described fall readily into two divisions (see Table, p. 307), ac-
cording to the nature of the cost at the proximal end of the
corallum. In one division the proximal end is smooth, that is
the coste on it are smooth and inconspicuous, or even hardly
discernible, and only come into evidence in the Trochoid shape-
stage ; inno stage do they appear granular. In the second division
the Peduncle or lowest Trochoid shape-stage is granular; the
granules may appear dispersed irregularly or may fall into definite
lines and tend to form irregular ridges, and in all cases they have
become the costal ornament by the time the Trochoid shape-stage
is reached. The first division contains the species P. centralis
(Mantell) and P. serpentina Edwards & Haime. The described
English species of the second division are P. fittont Edwards &
Haime, P. granulata Duncan, P. gravest Edwards & Haime (Dun-
can’s interpretation), P. mantelli Edwards & Haime, P. cylindrica
Edwards & Haime, and P. monilis Duncan (as here interpreted).
At first sight it is difficult to see any connection between the
coste of the first and those of the second division. Buta detailed
examination of the ontogenetic development of the cost renders
it likely that the primitive costal stages of the second division
represent the adult costal stages of the first division. At any
rate, non-granular stages precede the granular in the second
* P. M. Duncan, 1868, “ British Fossil Corals,” Mon. Pal. Soc. p. 58, pl. xvi.
figs. 5-15. Compare, for instance, fig. 6 (B.M. no. R. 120438) with fig. 14 (B.M.
no. R. 12051).
288 MR, .W. D. LANG ON GROWTH-STAGES | Mar. 2,
division. In each species the coste, in passing from the proximal
to the distal end of the corallum, exhibit very definite growth-
stages; and these costal stages are dependent on changes in two
characters—the general shape of the costa and its ornamentation.
The changes in the general shape of the costa are from a broad,
low, indistinct, to a comparatively narrow, high, and clearly
marked costa, and a corresponding katagenetic phase. The kata-
genesis * is nearly always rapid compared with the anagenesis *,
and each is represented on the diagram, text-fig. 38, by a steep limb
of the wave-like progression. It appears that thrice over in the
series under consideration has this period of costal rise and fall been
repeated, and each time with a different type of ornamentation.
When present, the ornamentation of the costa is of two kinds.
In the most primitive stages the costs are plain, or very slightly
rough, and only change in height. ‘This period is therefore here
called the Plain Period (text-fig. 38). The first marked orna-
mentation is a longitudinal etching as if the costal surface had
been corroded. As this becomes more intense, the ridges tend to
mass together and fuse, until the highest development is a wavy
ridge down the middle of the costa. Correlated with this etched
ornamentation is a second period of costal rise, the broader lower
costa having the irregular etching, and the high narrow costa the
ridged or nainent : this is the Etched Period. The second and
more advanced type of ornament is the granule, and the Granular
Period corresponds with a third costal rise.
Nine main stages may thus be formulated in this series of
Corals, namely,a low, medium, and high stage in the Plain, Etched,
and Granular Periods. Katagenetic stages have not been con-
sidered, because as a rule they are passed through too quickly to
be appreciated. Of course, no one species of Parasmilia shows
all nine stages; in fact the first three are so condensed in P. cen-
tralis (Mantell), one of the more primitive species, that a Plain
Period would not have been suspected to have had a separate
existence from the Etched Period, had not the scheme suggested
it when the other stages were marshalled in order. A more
careful examination ot P. centralis (Mantell) then showed a
heightening of the costa (costal-stage III.) before the normal low
etched stage (costal-stage TV.), thins revealing a Plain Period,
characteristic of P. serpentina Kdwards & Haime, preceding the
characteristic Htched Period of P. centralis (see text-fig. 40, F, G,
p- 295). Such Tachygenesis fT is, however, to be looked for when
cases of individual development in other phyla are remembered.
Again, the greatest gap in the series is between P. centralis
~ (Mantell), the most advanced costal-stage of which is the high stage
of the Etched Period (costal-stage VI.), and P. fittont Edwards &
Haime, the greater extent of whose costz is in the low normal stage
* A, Hyatt, 1889, “Genesis of the Arietide,’ Smithsonian Contributions to
Knowledge, Washington, xxvi. Art. 2, pp. 71-74.
+ A. Hyatt, 1893, “ Bioplastology and the related branches of Biologic research,”
Proc. Boston Soc. Nat. Hist., vol. xxvi. p. 77.
289
IN BRITISH SPECIES OF CORALS,
of the Granular Period (costal-stage VII.). To detect and interpret
the stages of the earlier parts of P. fittoni before costal-stage VII.
1909.]
Text-fig. 38.
Plain Pertod. Llched Pervoad. Granular lero
———_— LL __,
Anagenelic Phase. alagert| A nagenelce Phase. payee) Arnagerelic etapa
Stagel. S ree Stagel. | SlagelV. Stage V. Stage WZ i Ww CSE EL Stage B
| 72 ie ae € Haume.
| ee oe 4 a Une | |
| PManleltt,Zolwards & Haiwme. ee
P Cravest, Ll. & Hatin. er
73 Granwulace, D2ARCA2. ee ee Si,
| | Saree | | | | l
|
(2 cenGrates Martel).
3 Ses iS Sess | P. serpentina ,Zduards & Haime.
_ os L
Diagram illustrating the progress of costal ornamentation in the English species of Parasmilia.
But unless it is first
From this it will be realised how
close observation.
lly the case has been stated.
diagrammatica
requires
290 MR. W. D. LANG ON GROWTH-STAGES [ Mar. 2
presented in its baldest and most diagrammatic aspect, the extra-
ordinary regularity in the genetic sequence of the costal character
may be lost sight of in the ‘greater or less amount of development
and condensation of the various characters, and in certain irregu-
larities in their actual sequence due to environmental and other
causes. One such case will be next considered.
III. RecaprruLation oF GRowTH-STAGES AT REJUVENESCENCE.
The life-history of the costa in an actual specimen of each of
the described forms of English Chalk Purasmilia will be given
later on, that of the type-specimen when available, and in the
cases where this is not in the British Museum, one of Duncan’s
figured specimens if possible. In two cases specimens are de-
scribed which have no historical value in point of view of identi-
fication. But before describing these actual cases, an important
modification in the regular sequence of growth-stages must be
noted, Periodically in the growth of the corallum constrictions
arise, marking points in the life of the coral when fresh growth
took ‘place after a period of rest. Such revivals of erowth are
known as Rejuvenescence*, a term very happily introduced,
because the coral literally renews its youth at these points, and
expresses the renewal by recapitulating earlier growth-stages in a
condensed form; at least this is so in the costal characters of
Parasmiliay. A glance at the actual costal life-histories given
hereafter (see este: figs. 39-46) will show the extent of this
recapitulation.
There is no reason for supposing that the recapitulation of
younger stages at rejuvenescence is confined to the two characters
observed, nor only to the genus in question; rather, it probably
extends to the internal structure and is of general application.
The anomaly then in Caninia cornucopie Michelin, described by
Carruthers and mentioned early in this paper, may be explicable
by this phenomenon. The coral had developed as far as the
nystiana-stage, had next entered a period of rest, and on re-
Juvenescence, with renewed growth, had begun in the dumonti-
stage, recapitulating its grow uh from that stage to the nystiana-
stage again, and then proceeding, perhaps through further
interruptions of rejuvenescence and consequent recapitulations,
to the edwardsiana-stage.
TV. Bernarp’s THeory or THE NATURE OF REJUVENESCENCE
It is difficult to dissociate rejuvenescence from branching when
viewed in connection with recapitulation of earlier stages. Robert
« Verjtingung-process, ” Milaschevitch, 1876, Palaontographica, vol. xxi. p. 194-5.
3 ates Tomes, 1882, Quart. Journ. Geol. Soc. » p. 409; 1884, op. cit.
p. 363; 1886, Geol. Mag., p. 394; 1888, op. cit. p. 208 ; 1899, op. cit. p- 305.
iP It is the phenomenon here described that Bernard refers to iia a footnote,
British Museum Catalogue of Madreporarian Corals, 1906, vol. vi. p.
1909. ] IN BRITISH SPECIES OF CORALS. 291
Tracey Jackson * has shown how generaliy this takes place among
plants and is exhibited by the leaves. In many plants these show
progressive development when traced from the proximal to the
distal end of the branch, and the stage at the proximal end is an
earlier one than that of the leaf subtending that branch. A similar
case has been suggested among Polyzoa, concerning the change
in general shape of the zocecium‘t. Reduced to its simplest
expression, branching in corals is seen to become simple fission
in the vertical plane, exhibited by such a form as Zhecosmilia
[Chorisastrea| rugosa (Tomes) non Laubet, which is only a
Montlivaltia which has taken to division by fission; the Montli-
valtia-stage of this coral is very near J/. painswicki Duncan §,
which comes from the same horizon and district—the Aalenian
of the Cotteswolds. Another case is the specimen figured by
Duncan as Thecosmilia obtusa|| (d’Orbigny), which is only a
Montlivaltia fairfordensis 4 Tomes, which has begun to divide by
fission: both come from the Bathonian clay of Fairford, Gloucester-
shire, and these forms as well as fully divided ones are common
enough among the material from there. The examples quoted
are exhibited in the British Museum, where all the specimens
mentioned in this paper, unless otherwise stated, are to be seen.
In some Ceelenterates fission normally occurs in the horizontal
plane, resulting in the well-known phenomenon of strobilization.
It is obvious that if horizontal fission were to occur in a coral, the
upper product would obliterate the lower, since, having no means
of locomotion, it would continue to grow in place and would
secrete its skeleton on the top of the old **. That this new skeleton
would be in strict continuity with the old is rendered likely by
the soft nature of the secretory surface, which by its own weight
would be moulded on the underlying old skeleton, and, when
secreted, the new skeleton would reproduce the shape of, and so
fit, the old. An interesting case showing this is afforded by a
specimen exhibited in the British Museum (No. R.9148) of the
Montlivaltia-stage (M. fairfordensis Tomes) of Thecosmilia obtusa
(d’Orbigny), growing on an oyster with a ribbed shell. The ribs
of the oyster-shell are continuous with the coste of the coral
which has settled on it.
* R.T. Jackson, 1899, “ Localised Stages in Development,’ Mem. Boston Soc.
Nat. Hist. vol. v. no. 4, pp. 92, 131.
+ W. D. Lang, 1905, Geol. Mag. pp. 259, 260.
{ R. F. Tomes, 1882, Quart. Journ. Geol. Soc. p. 428; B.M. no. R. 10847,
§ P. M. Duncan, 1872, Mon. Pal. Soc., Part ui. p. 17, pl. 1. fig. 12; B.M. no.
R. 2305.
|| PB. M. Duncan, 1872, Mon. Pal. Soc., Part i. p. 14, pl. i. figs. 1-4; B.M. no.
R. 8455.
q R. F. Tomes, 1883, Quart. Journ. Geol. Soc. pp. 181, 182, pl. vii. fig. 21; B.M.
no. R. 8469.
** H. M. Bernard, 1906, British Museum Catalogue of Madreporarian Corals,
vol. vi. p. 22.
bo
Wo)
i)
MR. W. D. LANG ON GROWTH-STAGES [ Mar. 2,
V. REJUVENESCENCE AND TABULA- FORMATION.
Bernard *, who put forward this explanation of Rejuvenescence,
goes further and suggests that tabula-formation has a similar
origin. This is in direct contradiction to the generally accepted
idea that tabula-formation is only a more complete form of dis-
sepiment-formation ; that whereas the latter only expresses a
frequent and partial recession of the soft parts from the lower
portion of the skeleton as the coral grew upwards, so tabulee
express a periodic upward retreat of the soft parts as a whole.
This may well be the origin of tabule (dissepimental tabula) in post-
Rugose Madreporaria, the apparent tabule of which in many cases
seem to be modified dissepiments and often are seen to accompany
the reduction of septa. This is so in a series of Bathonian species
placed in various genera, but all allied to Cyathophora and Crypto-
cenia, of which a series is exhibited in the British Museum.
These show the tendency to lose their septa and convert the
interseptal dissepiments into tabule. Intermediate forms occur
between Cyathophora prattt Kdwards & Haime, with well developed
septa and few large dissepiments which tend to correspond with
similar ones on the other side of the corallite, and Astrea bowrgets
Defrance, in which the septa are only ridges and there are clearly
marked tabule. Holocystis elegans (Fitton), from the Lower
Greensand, has tabule and reduced septa, and the same is true of
the Paleozoic Tabulate corals. That tabula-formation of this
origin may have followed and replaced their formation from a
transverse fission 1s possible; but the two methods are essentially
distinct and mutually exclusive. The recapitulation of earher
stages might be used asa test to determine which cause is operative
in any case, for there is no reason to suppose that it would occur
merely on the upward movement of the coral in its skeleton; while
it has been shown to occur as an accompaniment of rejuvenescence.
VI. AstoGENy AND Hisronysis.
Before leaving the subject of rejuvenescence connected with
fission in a horizontal plane, it is interesting to note that Bernard
claims a similar behaviour in the coral colony to that m the indi-
vidual. Colonies of corallites, he says (that is, whole corolla),
undergo rejuvenescence, forming the well-known masses of coral
in cake-shaped pieces piled one on the other (metamerically seg-
mented, Bernard suggests), or else branched in regular order of
repetition. And he mentions an observation by Duerden { of a
living form which suggested that at the periods of rejuvenescence
a general histolysis took place in the coral mass, representing col-
lectively the fission of each individual. Again, the Polyzoa afford
* H. M. Bernard, 1906, Joc. cit. p. 23.
+ H. M. Bernard, 1906, British Museum Catalogue of Madreporarian Corals,
vol. vi. top of p. 24.
~ Duerden, 1904, “The Coral Siderastrea,’ Washington, Carnegie Institution
( tide Bernard).
1909. ] IN BRITISH SPECIES OF CORALS. 293
a parallel example of the behaviour of the colony as an individual.
Jn these animals the whole zoarium (colony) undergoes growth-
stages just as does the individual (zoecium). This was first
pointed out in Polyzoa by Cumings*, and he termed the colonial
developmental history Astogeny, as contrasted with the individual
Ontogeny.
VII. Inpivipuat Cosrat Lire-Hisrories of THE DESCRIBED
Species oF Encuish Parasurcrd. (Text-figs. 39-46).
[ Note.—The letters B.M. in the following descriptions mean “British Museum.”
The stages in the life-history of the individual are lettered thus : A, B, C,..&c.;
the costal-stages of the whole series; {., II., IIL, &c.]
1. PARASMILIA SERPENTINA Edwards & Haime. (Text-fig. 39.)
Parasmilia (?) serpentina Kdwards & Haime, 1850, “ British
Fossil Corals,” Mon. Pal. Soc. p. 51, pl. vii. figs. 3, 3a, 3 6.
Type-specimen, B.M. no. 48414.
Text-fig. 39.
PARAS/UILIA SERPENTINA,
Ldwards & Haime.
E* CYLINDRICAL STAGE.
REJUVENES CENCE.
CYLINDRICAL STAGE.
REJUVENESCEN CE «
CYLINDRICAL STAGE .
TROCHOID STAGE.
<< REJUVENES CENCE.
LTROCHO/D STAGE.
% a
= REIWVENES CENCE ,
Diy.
PEDUNCLE STAGE.
OY
Costal stages in P. serpentina Edwards & Haime.
Type-specimen, B.M. no. 48414. x 24.
* E. R. Cumings, 1904, American Journal of Science, series 4, vol. xvii. p. 49.
294 MR. W. D. LANG ON GROWTH-STAGES [ Mar. 2,
In this specimen the Peduncle, Trochoid, and Cylindrical shape-
stages are all long and pass gradually from one to another, giving
the corallum a regularly tapering shape. A rejuvenescence very
early in life makes a convenient demarcation between the Peduncle
and Trochoid shape-stage. The Peduncle being smooth for the first
quarter of its course, there is a Precostal stage (text-fig. 39, A).
Low plain cost (text-fig. 39, B, costal-stage I.) succeed, and
reach nearly to the first rejuvenescence ; but just before this the
costze become higher (text-fig. 39, C’, costal-stage II.), and are
almost ridged (text-fig. 39, D*, costal-stage III.) when reju-
venescence occurs. On resumption of growth the coste are in
costal-stage IT. (text-fig. 39, C*), but rapidly pass again to costal-
stage ITT. (text-fig. 39, D’), which is the characteristic stage of
the corallum as a whole, and continues to the distal side of
the third rejuvenescence ; that is, at the second and third re-
juvenescences the recapitulation is so Slight that costal-stage IT.
is not repeated. At about halfway between the third and four th
rejuvenescences there is a sharp katagenesis (text-fig. 39, E’),
marked by a lowering of the costa and “the disappearance of the
ridge. The cost then take on an etched ornamentation (text-
fig. 39, F", costal-stage IV.), but are thrown back again to costal-
stage ITT. (text-fig. 39, D*) after the fourth rejuvenescence. The
katagenesis and advance to costal-stage TV. is again repeated
(text-fig. 39, EH’, EF’), and then the corallum terminates. The
Trochoid shape-stage passes insensibly into the Cylindrical between
rejuvenescences two and three.
2. PARASMILIA CENTRALIS (Mantell). (Text-fig. 40.)
Madrepora centralis Mantell, 1822, “ Fossils of the South
Downs,” pp. 159-160, pl. xvi. fig. 4.
Type-specimen, B.M. no, 5462.
On the expanded base of the type-specimen there is no appear-
ance of costze (text-fig. 40, A). This Precostal stage is followed
by very faint, low, plain coste on the Peduncle and lowest Trochoid
shape-stages (text-fig. 40, B, costal-stage I.). By the mid-
Trochoid shape-stage the cost, though still plain (or very faintly
roughened), have become more prominent and higher (text-
fig. 40, C, D, costal-stages II. & IIT.). They then undergo a rapid
katagenesis and appear low again, but this time markedly etched
(text-fig. 40, E, costal-stage [V.). Shortly before the first re-
juvenescence the cost are heightened (text-fig. 40, EF’, costal-
stage V.) and become ridged (text- fig. 40, G’, costal stage VI).
Between the first and second rejuvenescences the shape passes
from Trochoid to Cylindrical, and costal-stages V. and VI. are
repeated (text-fig. 40, F’, G*); and though ‘there is no definite
rejuvenescence after the second, alternations of costal-stages V.
and VI. occur (text-fig. 40, F", G") and show periods of growth
alternating with fleeting rest-periods.
1909. | IN BRITISH SPECIES OF CORALS. 295
Text-fig. 40.
_ FPARASITULIA CENTRALIS, Mertz.
CYLINDRICAL STAGE.
— REIUVENESCENCE.
, CYLINDRICAL
\ STAGE.
G \
FE b | TROCHOlD STAGE.
Ee = S —PEJUVENESCENCE.
E. ul "
JE.
ya} TROCHOID
C: STAGE.
Bi:
B. PEDUNCLE STAGE .
AL
lam
Es
Costal stages in P. centralis Mantell.
Type-specimen, B.M. no. 5462. X 2.
3. ParasMILIA Firront Edwards & Haime. (Text-fig. 41.)
Parasmilia fittoni Edwards & Haime, 1850, “ British Fossil
Corals,” Mon. Pal. Soc. p. 50, pl. 1x. fig. 2 (non figs. 2 a, 2 6).
Type-specimen, B.M. no. 48412.
In their Monograph on British Fossil Corals, Edwards &
Haime state that Parasmilia fittont is in the collections of the
Geological Society, of the Geological Survey, and of the Museum at
Paris. This species 1s not mentioned in Blake’s and Sherborn’s
list of types in the collection of the Geological Society ; nor do the
Geological Survey possess the types; while at Paris there is the
specimen figured on pl. ix. figs. 2a, 2b; but) mot tig: 25) the
British Museum specimen No. 48412 agrees with Edwards &
Haime’s description and exactly corresponds with fig. 2 (reversed
in printing so that the figure is a mirror-reflection of the specimen)
and evidently is one of Edwards & Haime’s two types. It follows
then that there are two syntypes of this species, B.M. no. 48412,
pl. ix. fig. 2 of Edwards & Haime, and the Paris specimen figured
on pl. ix. figs. 2 a, 2 6; nor is it certain without examination of
296 MR. W. D. LANG ON GROWTH-STAGES [ Mar. 2,
the Paris example that the two specimens are specifically identical.
‘To save further confusion the specimen at hand, B.M. no, 48412,
is considered here as the holotype of P. fittoni; and the specific
determination of the Paris specimen must be settled when an
opportunity of examining it occurs. I wish to express my thanks
to Dr. Kitchin, of the Museum of Practical Geology, and to
Professor Boule, of the Muséum d’Histoire Naturelle, Paris, for
their kindness in supplying me with information in this matter.
Text-fig. 41.
PARASM/LIA FIT TON Edwards
& Haime.
D*
RESUVENESCENCE.
— REJUVENESCENCE.
g
5
s
REJUVENESCENCE.
J
[ee REJUVENESENCE.
O REJUVENESCENMCE,
Ss om RESUVENESCENTCE.
eee Gin
Sn / PEDUNCLE
STAGE.
Costal stages in P. fittoni Edwards & Haime.
Type-specimen, B.M. no. 48412. X 14 (Peduncle restored.)
In the forms hitherto considered, the costal growth-stages have
been easy to follow, because the ornamentation of the costa has
regularly followed the changes in its general shape. Six stages
have been demonstrated covering two periods—the Plain and
Ktched Periods; so that the stages have been termed Low Plain,
Medium Plain, High Plain, Low Etched, Medium Etched, and
High Etched. With P. fittoni a new type of ornamentation is
introduced—the granular ornament, and with it a new period of
costal elevation, But so dominant is the granular ornamentation
in P. fittoni, that instead of first appearing with a low wide costa,
1909..] IN BRITISH SPECIES OF CORALS. 297
it comes in while the costs are still high after costal-stage VI.
(the High Etched) and katagenetic as far as height goes from
costal-stage VI. towards the low costa of costal-stage VII. So
far the katagenetic phases of each period have been inconsiderable,
but the fall in height from costal-stage VI. to costal-stage VII.
occupies much of the Trochoid shape-stage in the type of P. fittoni.
In this specimen the peduncle is absent and the costa on first
appearing are in costal-stage VI., the High Etched stage. ‘The
earlier stages are seen in the B.M. specimen No. R. 6632, figured
as Monocarya centralis Mantell by Lonsdale, 1850, in Dixon’s
‘ Geology of Sussex,’ pl. xviii. fig. 4. This specimen, unfortunately
damaged in the Trochoid shape-stage, exhibits the earlier and
later parts and agrees very closely with the type specimen of
P. fittoni. The costee when first visible are low and wide, but
rapidly become narrow and high with an ornamentation consisting
of massed granules (text-fig. 41, A, costal-stages [V.—V.-VI.). As
far as this first high stage the ornamentation appears as merely
a roughness and is considered as indicating the Etched Period.
The Peduncle shape-stage may be looked upon as including the
Plain and Etched Periods of costal development reduced by
Tachygenesis to a mere rise in the costa accompanied by a
roughening of the surface. Whether this is the true explanation
or not, it is a fact that there is a period of costal elevation which
has declined before the appearance of the typical granular
ornament of P. fittons.
On returning to the type-specimen B.M. no, 48412, it is found
that the peduncle is broken off, so that the corallum begins in
the lowest Trochoid shape-stage. The costs when first clearly seen
are high and narrow (text-fig. 41, B), but rapidly widen and become
lower (C) until, by the middle of the Trochoid shape-stage,
they are wide and low and by this time have assumed the typical
ornamentation of the species, which is very small granules, in-
dividually separate (text-fig. 41, D', costal stage VII.). At first,
with the high cost, the ornament appears asa rough ridge, which,
ag the costa widens, becomes lower, resolves into massed granules
and finally (costal-stage VII.) into granules individually separate.
This katagenesis suggests a previous granular anagenesis, but the
stages on the peduncle do not suggest this; they are etched rather
than granular, and it is more probable that the granules of costal-
stage VII. have appeared earlier than thei appropriately shaped
costa and have become massed and piled in consequence of the
shape of the costa.
At about the mid-Trochoid shape-stage, before the coste have
reached costal-stage VII., the first rejuvenescence occurs. It
does not, however, interrupt the progress of the costal stages
by causing recapitulation. This and the next rejuvenescence are
partial and only affect the convex side of the corallum, which, as
a whole, is considerably bent.
Before the second (partial) rejuvenescence, the costa become
narrower and the granules begin to mass together (text-fig. 41, KH’,
Proc. Zoou., Soc.—1909, No. XX, 20
298 MR. W. D, LANG ON GROWTH-STAGES [Mar. 2,
costal-stage VIIT.). On resumption of growth costal-stage VII.
reappears (text-fig. 41, D*) succeeded by costal-stage VIII. before the
third rejuvenescence (text-fig. 41, E*). This sequence is repeated
before the fourth and fifth rejuvenescences (text-fig. 41, D*®, D‘-
E’, E’). The fifth and sixth rejuvenescences are compound ones,
several occurring in both cases in quick succession. In both,
too, the coste reach a more elevated condition and the massed
granules tend to form a ridge before rejuvenescence occurs (text-
fig. 41, F’, F’). This stage is not advanced enough to be con-
sidered costal-stage [X.,and does not recur in the corallum of this
specimen though there is a seventh rejuvenescence. The fifth,
sixth, and seventh rejuvenescences throw the cost back to costal-
stage VII
4, PARASMILIA GRANULATA Duncan. (Text-fig. 42.)
Parasmilia granulata Duncan, 1869, “ British Fossil Corals,”
Mon. Pal. Soc. p. 13, pl. vi. fig. 5.
Monocarya centralis Lonsdale, 1850, in Dixon’s ‘ Geology of
Sussex,’ pl. xviii. fig. 2.
Text-fig. 42.
PARASMILIA
GRANULATA , DUNCAN.
CYLINDRICAL STAGE.
—REJUVENESCENCE.
ae T-ROCHOI/D STAGE.
4.
Gs __ REJUVENESCENCE .
Pp
= — REVUVENESCENCE.-
= TROCHO/D STAGE .
tis — REJUVENESCENCE.
E z
é REJUVENESCENCE .
é/
TROCHOID STAGE.
Dd.
[EHO Grr? 2s 4 nee
pt PEDUNCLE STAGE.
A UL
Costal stages in P. granulata Duncan.
Type-specimen, B.M. no. R. 6612. &X 2.
909.) IN BRITISH SPECIES OF CORALS. 299
Type-specimen, B.M. no. R. 6612. (The specimen figured in
both the Plates referred to above.)
P. granulata very closely resembles P. jittont Edwards & Haime,
but costal-stage VII., the typical costal-stage of P. fittoni, is very
reduced and replaced in importance by costal-stage VIIL. ; also
costal-stage IX. is more nearly attamed in P. granulata ; other-
wise the costal-stages closely correspond. But the katagenesis in
the height of costa from costal-stage VI. to costal-stage VII. is
even more prolonged than in P. fittoni, and the first appearance
of granular ornament pushed back to costal-stage VI. ‘The
individual stages in P. granulata (text-fig. 42, A, B, C... &e.)
will be described in order, so that the points above mentioned
may be made clear.
A=Costal-stage 1V. The Primary coste are double, or longi-
tudinally split. Coste low; ornament etched.
B=Costal-stage V. Similar to stage A, but coste of medium
height.
C=Costal-stage VI. Primary cost single and so for the
remaining stages. Coste high, narrow; massed granular
ornamentation.
D=Costal-stage VII. for the ornament which is of individually
separate granules, though not so separate as in P. fittont ;
and katagenetic for the height of costa from costal-stage VI.
to costal-stage VII.
E'=Costal-stage VIII. for the ornament which is of massed
granules, and katagenetic for the height of costa from
costal-stage VI. to costal-stage VII.
First Rejuvenescence.
E’. Similar to EK’.
Second Rejuvenescence.
F'=Costal-stage VII. for both height of costa and qeraent
z. @. a br onal low costa and gr antes individually separate.
G'=Costal-stage VIII. Coste a medium height and breadth ;
granules massed together and tending ‘a form a Oeatienl
ridge.
Third Rejuvenescence.
FE? & G®. A repetition of stages FP’ & G’.
Fourth Rejuvenescence.
F* & G*. A repetition of stages F” & EF’, G’ & G’.
H=Stage between costal- stage VII. and costal-stage IX,
Closes high with thin wavy ridge, but massed evanules are
still deat:
Fifth Rejuvenescence. Trochoid shape ..age ends and Cylin-
drical shape-stage begins.
F & G*. A repetition of stages F’ &ec., G* &e.
20*
300 MR. W. D. LANG ON GROWTH-STAGES [ Mar. 2,
5. PARASMILIA GRAVESI Edwards & Haime. (Text-fig. 43.)
Parasmilia centralis (Mantell), sub-species gravesana Edwards
& Haime, Duncan, 1869, “ British Fossil Corals,” Mon. Pal.
Soc. p. U2 spliwva tie .08
In the absence of the type-specimen it is necessary to take
Duncan’s interpretation of this species, and the specimen he
figured, B.M. no. R. 6616, is here described as P. gravest Kdwards
& Haime, on the assumption that Duncan was correct in his
determination.
Text-fig. 43.
PARAS/ULIA GRA VEST fiwards & Haime.
MW.
2 10 G?- = 5
/o Qa
Be
&.
fe
ay ee CYLINDRICAL STAGE .
E3. Y __ REJUVENESCENCES .
G! TROCHOID STAGE .
: — RES UVENES CENCE
E. ; TROCHOLD STAGE.
ES
SRN Mant hy
a ; EESERCE
TROCHO/D
STAGE.
D. |
Ge
e-em Wa EDUNCLE STAGE.
ae WIE
Costal stages in P. gravesi Edwards & Haime.
Duncan’s figured specimen, B.M. no. R. 6616. X 2.
Specimen No. R. 6616 carries the modifications of P. fittoni
exhibited in P. granulata still further. Costal-stage VII. (when
once established for both height and ornament) is hurried through
and hardly appears ; costal-stage VIII. is important ; and costal-
stage IX. is actually attained. There is a very prolonged kata-
genesis in the height of the costa from costal-stage VI. to costal-
1909.] IN BRITISH SPECIES OF CORALS. 301
stage VII.; and the first appearance of granular ornament: is
pushed back to costal-stage V.
At the extreme edge of the peduncle the coste are low and
have an etched ornament (text-fig. 43, A, costal-stage IV.). On
the rest of the peduncle they are higher (text-fig. 43, B, costal-
stage V.), but the ornamentation is resolving into granules.
On the lowest Trochoid part of the corallum the costs are high
(text-fig. 43, C, costal-stage VI.) and the ornamentation is of
individually separate granules and is the ornament of costal-
stage VII. pushed backwards. The primary coste are forked at
this stage, the branches crossing neighbouring cost (cf. stages
A and B of P. granulata). From this point the coste decrease in
height and increase in breadth (text-fig. 43, D, katagenesis from
costal-stage VI. to VII.) until past the mid-Trochoid shape-stage ;
here the coste are broad and low with individually separate
granules (text-fig. 43, E*, costal-stage VII.), but immediately these
mass and the costa becomes narrower and higher (text-fig. 43, F’,
costal-stage VIII.), and then rejuvenescence occurs. Between
this and the second rejuvenescence costal-stage VIII. (text-
fig. 43, F*) is dominant, but costal-stage VII. is just represented
(text-fig. 43, E*) and costal-stage IX. (text-fig. 43, G*) is just
attained. Rejuvenescences 3, 4, and 5 cause the repetition of
costal-stages VII. and VIII. (text-fig. 43, E®°, F*’) but not of
costal-stage IX., and costal-stage VII. is only just indicated.
The Cylindrical shape-stage has now been reached and a series of
very slight constrictions indicates rejuvenescences 6, 7, 8, and 9,
with a similar repetition of costal-stages VII. and VIII.; and
before rejuvenescence 7 costal-stage IX. is repeated (text-fig. 43,
KY’, F°°, G*). The repetition of costal-stages VII., VIII., and
IX. (text-fig. 43, E, F'°, G’) precedes rejuvenescence 10, which
is a large one, and the corallum ends in costal-stage VII. (text-
fig. 43, E'*) which is assumed when growth recommences,
6, PARASMILIA MANTELLI Edwards & Haime. (Text-fig. 44.)
Parasmilia mantelli Kdwards & Haime, 1850, “ British Fossil
Corals,” Mon. Pal. Soc. p. 49, pl. viii. figs. 2, 2 a.
Type-specimen, B.M. no, 48413.
P. fittoni Kdwards & Haime, P. granulata Duncan, and P. gravest
Edwards & Haime, form a group of closely allied species, pro-
gressing serially in costal development from a condition in which
costal-stage VII. is dominant to one in which costal-stage VIII. is
more important; and costal-stage IX. is hardly attained. More-
over, in this series there is always a long katagenesis after costal-
stage VI. and before costal-stage VII. is attained. The group of
P. mantelli Edwards & Haime, P. cylindrica Edwards & Haime,
and P. monilis Duncan (as here determined) presents a series
progressing from a condition with costal-stage VIII. to one with
costal-stage IX. dominant, and though costal-stagé VII. may be
302 MR. W. D. LANG ON GROWTH-STAGES [Mar. 2,
represented, it appears, when present, immediately on the
peduncle and without any preliminary long katagenesis from
costal-stage VI. Also intercostal dissepiments occur on the
three species of this group.
Text-fig. 44.
PARAS/MILIA MANTELLIT,
Eilwards & Hamre.
TROCH OID
STAGE.
—— RESUVENES CENCE
TROCHOID
STAGE.
Cc:
C. PEDUNCLE STAGE.
B.
vals
Costal stages in P. mantelli Edwards & Haime.
Type-specimen, B.M. no. 48413. x 3.
As soon as cost appear on the peduncle of P. mantelli, they
are low and ornamented with few large granules which tend to
mass together (text-fig. 44, B, costal-stage VII., A is the pre-
costal stage). The coste then rapidly rise in height, become
thinner and the granules become heaped in masses (text-fig. 44, C,
costal-stage VIII.). At about halfway in the Trochoid shape-
stage intercostal dissepiments occur; and from here to nearly
the top of the Trochoid shape-stage the granules tend to dis-
appear from the sides of the coste and to form a ridge along
their top (text-fig. 44, D', stage between costal-stages VIII.
and IX.). At the top of the Trochoid stage the coste have
become high and thin with a sharp wavy edge, and the granules
1909. ] IN BRITISH SPECIES OF CORALS. 303
have disappeared from their sides (text-fig. 44, E, costal-stage
IX.); intercostal dissepiments are still pr esent. Rejuvenese ence
now occurs and the corallum ends with resumption of growth
in costal-stage VIII. (text-fig. 44, D*), Probably the Gylindrie: al
shape-stage begins in this specimen after the first rejuvenescence.
7. PARASMILIA CyLINDRICA Edwards & Haime. (Text-fig. 45.)
Parasmilia cylindrica Edwards & Haime, 1850, “ British Fossil
Corals,” Mon. Pal. Soc. p. 50, pl. viii. fig. 5.
B.M. nos. R. 13889 & R. 6614.
Text-fig. 45,
PARAS/ILIA
CYL/NDRICA , Epwarns &HAIME.
Gir
Bis CYLINDRICAL
STAGE.
c* MANY
B REJUVENESCENCES.
C3
BX REJUVENESCENCE.
Cc -RESVENESCENCE. & 3
3B ~REJUVENESCENCE. bs .
a -Revenescence. ™ S
A S
5 ae
IG
Costal stages in P. cylindrica Hdwards & Haime.
B.M. nos. R. 18889 & R. 6614. > 1h.
The type-specimen of this species, from the Norwich Chalk, is
stated, by Edwards and Haime, to have been in the ‘ collection —
of Mr. J. 8. Bowerbank.” Though the Bowerbank Collection was
acquired by the British Museum in 1865, this specimen has not
been found. The species, however, is well-marked and easily
304 MR. W. D. LANG ON GROWTH-STAGES [ Mar. 2,
distinguished and specimens B.M. nos. R. 13889 and R. 6614,
from the Norwich Chalk, are here described. Probably these
two are the proximal and distal parts of one corallum; but as
they do not exactly fit together, it is not safe to assume that
they are one specimen, and the former is chosen as typical of
the species, because it shows the early as wellas the adult growth-
stages.
The cost are low when they first appear, with few, large granules
tending to mass (text-fig. 45 A’, costal-stage VII.), but soon
become thinner with a crest of massed granules (text-fig. 45, b’,
costal-stage VIIT.). The first rejuvenescence occurs very early in
the corallum and throws the coste back to stage VII. (text-
fig. 45, A*), whence they rapidly pass to stage VIII. (text-
fig. 45, B*), and again rapidly to stage IX. (text-fig. 45, C’) which
is characterised by high, thin, wavy coste with no granules and
with occasional intercostal dissepiments; the latter form an
irregular ring round the corallum at the second rejuvenescence,
which takes place low down in the Trochoid shape-stage. The
costal history need be described in detail no further, as there is
henceforth only a frequent alternation of costal-stages VIII. and
IX., the latter being predominant and the former tending to
disappear altogether in the Cylindrical shape-stage (text-
fig. 45, 1B Cay:
8. PARASMILIA MONILIS Duncan. (Text-fig. 46.)
Parasmilia monilis Dunean, 1869, ‘“ British Fossil Corals,”
Mon. Pal. Soe. p. 12, pl. v. figs. 4, 5, 6.
B.M. no. R. 6477.
The determination of this species cannot be regarded as settled
until the Type-specimen comes to light and has been examined, or
its loss certified and a new Type chosen. It is not easy from
Duncan’s figure to see if intercostal dissepiments are present ;
they certainly do not appear in enlarged figured pieces of coste
high on the corallum; on the peduncle, of which there is also an
enlarged drawing, they would not be expected.
A form represented by the B.M. specimen No. R. 6477 from
the zone of Belemnitella mucronata Schlotheim, of East Harnham,
Salisbury, from the Tomes Collection, is here described as P. monilis,
as it closely resembles Duncan’s figure. Small but numerous inter-
costal dissepiments are present in the distal part of the corallum,
and it is chiefly the presence of these that makes the determination
doubtful. Tomes’ specimen is one of a series described by him as
P. mantelli Edwards & Haime. There is also a specimen of
P. monilis as here defined, B.M. no. R. 6618, figured in 1850 by
Lonsdale in Dixon’s ‘Geology of Sussex,’ pl. xviii. fig. 9, as
Monocarya centralis (Mantell). This, however, has not been
chosen for description as the whole of the proximal end is gone,
and its exact horizon and locality are unknown.
P. monilis as here determined is remarkable for showing an
1909. ] IN BRITISH SPECIES OF CORALS. 305
even more advanced type of adult costa than P. cylindrica, in the
elimination of costal-stages VII. and VIII. from all but the most
proximal part of the corallum ; yet on the peduncle more primitive
stages are shown than even P. mantelli exhibits, and in this
P. monilis forms a link between P. gravest (as Duncan interpreted
it) and P. mantelli.
Text-fig. 46.
ae PARASPILIA
/STONILIS , DUNCAN.
— RESUVENESCENCE.
CYLINDRICAL STAGE.
— KESOVENESCEWCE .-
ae CYLINDRICAL
G. STAGE
Bs SRESUVENESCENCE
Gua
= TROCHOLD.
STAGE.
E..
/DY.
a. PEDUNCLE STAGE .
TU
Costal stages in P. monilis Duncan.
B.M. no. R. 6477. x3.
At the base of the peduncle the costz are low and rather rough
(text-fig. 46, A, costal-stage [V.), but rapidly heighten (text-fig. 46,
B& C, costal-stages V. & VI.). In this, the Etched Period, the
primary cost are longitudinally split (cf. P. granulata and P.
gravest). At the lowest part of the Trochoid shape-stage the costa
are low with few large granules (text-fig. 46, D, costal-stage VIL).
These immediately mass together and the costze rise in height (text-
fig. 46, H, costal-stage VIII.) and by the time the mid-Trochoid
shape-stage is reached, the granules have entirely vanished, the
coste are high with a thin, somewhat wavy ridge, and intercostal
dissepiments have appeared (text-fig. 46, F’, costal-stage [X.). A
306 MR, W. D. LANG ON GROWTH-STAGES [Mar. 2!
further stage (text-fig. 46, Gt), in which the cost are more wavy,
occurs before the end of the Trochoid shape-stage. At this point
the first rejuvenescence occurs, and on beginning growth again in
the Cylindrical shape-stage the corallum is in stage Ba (text-
fig. 46, F°), but rapidly passes to stage G (text-fig. 46, G*). After
the second rejuvenescence the wavy G stage seems to have become
permanently established, to the exclusion of stage F.
VIII. Summary.
Perhaps too much stress has been laid upon what, after all, are
only a few observations ; but the points introduced are of are
interest that it is hoped that they will be borne in mind and
tested when work is done on Corals, and not lost sight of in the
desire to describe new forms or to prove that too many already
have been described. Summarily the points touched on are
these :— Hereditary growth-stages exhibited in Rejuvenescence as
evidence that the latter is a form of Fission, and subject to the
same laws as other forms of Fission, such as s Branching ; ; views on
the formation of Dissepiments and Tabule and Bernard’s ideas
on these in connection with Fission; their equal application to
the colony as to the individual and a suggested relation with
Histolysis ; finally comparison with other widely different groups
of branching organisms— Plants and Polyzoa, that laws of
branching in one group may be compared with those in another
and any law common to all may be determined.
Nore :—Owing to the present inaccessibility of the collection
of British Chalk fossils formed by Dr. A. Rowe, it has not been
possible to test as fully as possible the stratigraphical value
of these observations; but thanks to the kindness of Messrs.
G. E. Dibley, C. P. Chatwin, and T. H. Withers, a certain amount
of zonally-collected material has been available for comparison
with the specimens here described. This, together with those of
the British Museum specimens whose zone is known, has made it
possible to draw up the following scheme of distribution in time.
It will be seen that the order of forms arranged according to this
distribution agrees on the whole with that deduced above from
their structure.
P. serpentina, and forms resembling it: zone of Holaster planws—
Micraster coranguinum.
P. centralis, and forms resembling it: zone of Holaster planus —
Micraster coranguinun.
P. fittoni, ;
P. granulata, and forms resembling them; zone of JV craster
P. gravesi, coranguinum.
P. mantelli.
e Hes iL Zone of Belemnitella mucronata.
. cylindrica. |
[8rd April, 1909.
Py Fg re by bo) Fy gg
1909.} IN BRITISH SPECIES OF CORALS. 307
TABLE CoRRELATING INDIVIDUAL Growrn-Sraces (A, B, C,....) witH
CostTaL-STAGES OF THE WuoLe Serres (I., II., IIL.,....).
N.B.—Where letters are not in continuous sequence (e.g. P. jfittoni, A, D, EB) the
missing letters represent katagenetic stages.
| {
|
Pre- Costal- Costal- Costal- Costal- ICostal- Costal- Costal- Costal- Costal-|
costal-, stage | stage | stage | stage | stage! stage stage stage stage |
| stage. 1a aay pice a i | Wa PP aVELep V2 UT VIII. | IX, |
. serpentina Edw. & H..... A Bae ice D ¥ | |
. centralis (Mantell) ......) A Baer © D Diol id) (Cet) |
ejuttont Bdw. & WH. .........| x x x x A | A A D E |
. gramidata Duncan ...... x x x x A || B Coe Bye Genk /
teraeni iw GM | x) x | x | x | A |B oe ian | F | G
. mantelli Edw. & H. DAM | | KI Ie oc Se wegen ns to
Neylind nico mbdwa ci RGus | xX! | x | Se SG is TAC rie lenin at
» mons, Duncan ....+.-. ere al eros x x A B C D E | F, G |
TABLE FOR DETERMINATION OF ENGLISH SPECIES OF Parasmilia.
A. Peduncle and lowest part of Trochoid shape-stage smooth or with
faint, plain cost; no granules; 1. e. costal-stage VIL. is not
attained.
( I. Cost over most of the corallum with sharp plain crest; i.e
H costal-stage ILI. is dominant ...... 1. Parasmilia serpentina Edwards
II. Coste over most of the corallum with rough, irregular mark- [& Haime.
ings, but not granular; 1. e. Héched Period is dominant.
2. Parasmilia centralis (Mantell).
B. Peduncle or lowest part of Trochoid shape-stage with granules ;
i. e. costal-stage VII. is attained.
( I. Granules are the costal ornament even in the most distal parts
of the corallum, though the granules may be massed to-
gether ; i.e. costal-stage LX. is not attained.
i In the parts of the corallum with granular cost# the
_granules are mostly individually separate and the costwz
low and broad ; i. e. costal-stage VII. is dominant.
| 3. Parasmilia fittoni Edwards &
6. In the parts of the corallum with granular coste, the [Haime.
granules mostly are more or less massed and the costz
comparatively high; 1. e. costal-stage V ILI. is dominant.
4. Parasmilia granulata Duncan.
| 1. At the distal end of the corallum the coste are thin, high
and sharp and without granules; 1. e. costal-stage LX. is
| attained.
(a. Granules vanish from coste only for short periods, before
a rejuvenescence; i.e. costal-stage LX. is hardly attained.
| ( No intercostal dissepiments; a long katagenesis be-
tween costal-stages VI. and VIL.
h 5. Parasmilia gravesi Edwards &
| 2. Numerous intercostal dissepiments ; costal-stage VII. | Haime.
tL immediately follows costal-stage VI.
6. Parasmilia mantelli Edwards &
b. Gains vanish from cost over long stretches of the (Haime.
corallum ; i. e. costal-stage LX. is easily attained.
(1. Coste on the peduncle on the whole are low and in-
conspicuous with many granules; i. e. costal-stage
| | | VIL. is well developed. Corallum is large, diameter
| | in cylindrical shape-stage 15-20 mm.
7. Parasmilia cylindrica Edwards &
| 2. Coste on the peduncle on the whole are high and { Haime.
granules not scattered; i. e. costal-stage VII. is
| | | poorly developed. Corallum is small, diameter in
cylindrical shape-stage 6-8 mm,
Gate 8. Parasmilia monilis Duncan.
L
308 MR. E. C. CHUBB ON TWO F@TAL LIONS. [Mar. 16,
March 16, 1909.
FREDERICK GILLETT, Esq., Vice-President, in the Chair.
The Secretary read the following report on the additions made
to the Society’s Menagerie during the month of February 1909 :—
The number of registered additions to the Society’s Menagerie
during the month of February last was 92. Of these, 50 were
acquired by presentation, 2 by purchase, 28 were received on
deposit, 2 in part exchange, and 10 were born in the Gardens.
The number of departures during the same period, by death and
removals, was 134.
Amongst the additions special attention may be directed to :—
One Common Tapir (Zapirus americanus) 8, from South
America, and one Malayan Tapir (Zapirus indicus) 2, from
Malacca, received in part exchange on February 2nd.
One American Bison (Bison americanus) 2, born in the Men-
agerie on February 27th.
The Secretary exhibited, on behalf of Mr. E. C. Chubb, F.Z.S.,
the skins and skulls of two feetal lions which had been presented
to the Rhodesia Museum, Bulawayo, by Mr. A. Giese. They had
been taken from a lioness which Mr. Giese shot last November at
Deka, about 50 miles south of the Victoria Falls.
These fcetuses showed very little pattern compared to that of
newly born cubs; the black markings in each example of the
former consisting only of a fairly well-defined median dorsal line,
a mottling on the head, some distinct spots on the outer sides of
the limbs, and a suspicion of rings on the tip of the tail.
The Secretary exhibited a photograph of a young American
Tapir, and called attention to the remarkable resemblance between
that and the young Malayan Tapir, a photograph of which was
reproduced in the Society’s ‘ Proceedings, 1908, p. 786. The
longitudinal light stripes on the flanks of the body, the spots on
the legs, and the white tips to the ears were present in both.
The Secretary called attention to an interesting paper by
C. Onelli in the ‘ Revista del Jardin Zoologico de Buenos
Aires,’ 1908, p. 207, in which the author described a general
correspondence between the number of vertebrae and the number
of stripes or rows of spots in many mammals.
The Secretary exhibited a photograph of a small herd of
Mountain Zebras (Equus zebra) in the possession of a dealer at
Port Elizabeth, South Africa.
PZ.S. 1909. Pl. XXXV.
#.Wilson, Gambridge.
GONIODES TETRAONIS.
P.Z.S. 1909. Pl. XXXVI.
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GONIODES TETRAONIS.
PZ.S.1909. Pl XXXVII.
H.Wils on, Cambridge.
TETRAONIS.
GONIODES
PZ_S. 1909) Pl XXXVile
1 mac.
E.Wilson,Cambridge.
GONIODES TETRAONIS.
PZ.S. 1909. Pl. XXXIX.
E. Wilson, Cambridge.
GONIODES TETRAONIS.
BAS, WOR) IPL IAL.
E. Wilson, Cambridge.
STRAONIS anpD MENOPON PALLESCENS.
————
SSS SSS)
PZ.S. 1909: PLXET.
E.Wilson ,Cambridg e.
GONIODES anpd NIRMUS.
PZ.S. 1909. Pl. XLII.
E. Wilson, Cambridge.
NIRMUS CAMERATUS.
PASS) USVONS). IL CLA
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ORNITHOMYIA LAGOPODIS.
E.Wilson, Cambridge.
P.Z.S:. WOSPEI Sai
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E.Wilson, Cambridge.
ORNITHOMYIA LAGOPODIS.
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SCATOPHAGA STERCORARIA
PZ.S: 190932 anvle
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7
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SCATOPHAGA STERCORARIA.
P Zuo. 19097 Plainville
E Wilson,Gambridge.
CERATOPHYLLUS GALLINULAE.
1909.] ON ECTOPARASITES OF THE RED GROUSE. 309
The Secretary also exhibited a photograph of a female Giraffe
captured in the West Soudan, east of Timbuctoo, showing a
general resemblance with Giraffa camelopardalis typica.
Dr. F. A. Bather, M.A., F.Z.S., exhibited a fossil Echinoid,
Scutellina patella, from the Hocene? (Barwonian) of Nelson,
Glenelg River, Victoria, Australia, showing a marsupium for the
young, as described by T. 8S. Hall (Proc. Roy. Soc. Victoria, n. s.
xx. p. 140, 1908).
The following papers were read :—
1. The Ectoparasites of the Red Grouse (Lagopus scoticus).
By A Ee SaipnEy, MAC) Elon DSey ER: S.,) Be ZS.
Fellow and Tutor of Christ’s College, Cambridge, and
Reader in Zoology in the University.
[Received November 28, 1908. ]
(Plates XXXV.—XLVITI.*)
I have made a careful examination of all the animals I can find
recorded both from the outside and from the inside of the
body of the grouse, with the exception of the blood parasites
Leucocytozoon lovati Seligm. & Samb.7, the Protozoa found by
Dr. Fantham, and the Microfilaria described by Sambon. Besides
the Protozoa found by Sambon, Dr. Fantham has found a species
of Spirochete in the blood prepared from smears taken from
the heart and liver. Probably this is a new species, as it is
somewhat short and thick. It was only seen on two occasions.
A second Spirochete was found in the intestine. It was seen
alive and was not numerous. It is possibly the so-called
Trypanosoma eberthi (Kent). Oocysts of Coccidium avium are
common in the intestine and rectum. Gregarine spores, probably
of Monocystis, are sometimes seen in the intestine; MJ/onocystis
spores having been recorded hy Pfeiffer from the intestines of birds.
A Heemosporidian occurs in the erythrocytes, but was seen only
in the younger stages. Theyare rare. An Amceba is found both
in the rectal contents and in the droppings.
The animals which live within the grouse constitute the Entozoa
or Entoparasites, those that live on the body form the Ectozoa or
Ectoparasites, and with these latter this paper deals.
From the point of view of the Grouse Disease Inquiry the
attention paid to the ectoparasites may seem superfluous, but
many of the internal parasites and all the tape-worms pass
through a second host. For example, the tape-worms which
live in the alimentary canal of the grouse pass their younger or
* Wor explanation of the Plates, see p. 333.
+ ‘The Lancet,’ 21 September, 1907, p. 829.
9
310 DR. A. E. SHIPLEY ON [ Mar. 16,
larval stages in the body of some lower animal. This lower
animal, presumably an insect or a molluse or a spider, must be
eaten by a grouse and the larval tape-worm must be set free
before the latter can grow up into the adult tape-worm which we
find in the intestine of the grouse. In searching for this second
host it was natural to begin with the ectoparasites, which one
would imagine were continually being snapped up by the bird.
We have, however, up till now completely failed to find any
cestode-larve in the grouse-fly or in the numerous ‘“ biting-lice ”
or “bird-lee” (Mallophaga) which abound on the skin and
amongst the feathers of the grouse; and, what is still more
significant and still more remarkable, we have, in the hundreds
of crop-contents which we have examined, never found one of
these insects in the grouse’s food.
This report is based in the main on my own observations, but
some of the facts recorded were first observed by Dr. E. A. Wilson,
and some by Mr. J. C. F. Fryer, of Caius College, Cambridge. In
fact, in looking back over the work | find it difficult to disentangle
the precise share each of us had in it. One thing, however, is
clear. I am indebted to Dr. Wilson for a very large proportion
of the drawings which have been reproduced in the Plates at the
end of this paper, and I am also indebted to him for lightening
many pleasant hours spent, not on the open, breezy heather of
the Scottish moors, but in the stuffy laboratory we were wont
to improvise in the back premises of many a Scottish inn.
To Mr. Edwin Wilson, of Cambridge, a word of thanks is also
due for the accuracy and skill with which he has depicted the
Grouse-fly and the Grouse-flea.
ECTOPARASITES.
INSECTA.
A. MatropHaca.—Bird-lice or Biting-lice.
Gi.) Fam. Philopteride.
I.—G@oNIoDES TETRAONIS Denny.
Tn his ‘ Monographia Anoplurorum Britannie,’ Denny * describes
and figures this species, which he calls the “‘ Louse of the Black
and Red Grouse.” He states that it is ‘‘common upon both the
Black and Red Grouse” (Lagopus tetrix and L. scoticus). ‘‘ Upon
the Willow or Hazel Grouse (Lagopus salicett) I find a similar but
distinct species, rather broader in the abdomen, and of much
darker colour.” Denny describes several species of the same
genus which infest other game-birds.
* Published by H. G. Bohn, London, 1842, p. 161, pl. xiii. fig. 3.
9
1909.] ECTOPARASITES OF THE RED GROUSE. dll
Giebel* gives the name (loniodes heteroceros Nitzsch as a
synonym of G. tetraonis, and in his large monograph on ‘ Les
Pédiculines,’ Piaget T uses the former name without any reference
to Denny's. The name G. heteroceros also appears in Giebel’s
article £ on the Epizoa of the Halle Museum, published in
1866, but only the name. In his article on “ "Parasiten ” in
von Middendorf’s ‘Reise in den Aussersten Norden und Osten
Sibiriens,’ Grube attributes certain bird-lice taken from Lagopus
albus, the Willow Grouse, and from Lagopus alpinus, the
Ptarmigan, to the species Goniodes tetraonis Denny; but Piaget
points out certain differences, and seems to consider that a new
species might have been described from these specimens.
Andrew Murray, in his book on ‘ Economic Entomology’ §,
writing of Goniodes tetraonis, says:—‘‘ This is the insect which
sometimes, especially in the bad seasons, does so much harm to
the young grouse when they are feeble and unhealthy.”
It is the commonest of the insects which infest the skin of
grouse, crawling about amongst the base of the feathers and on
the vane of the feathers themselves. It occurs more commonly
than Nirmus cameratus, which is often associated with it. It is
comparatively rare to find a bird free from these ‘“ biting-lice,”
but perhaps 10 per cent. is about a fair estimate of the number
of uninfested grouse. The number on each bird is to some extent
an inverse measure of their health. Careful search will discover
but two or three on a healthy grouse, but on a “ piner ” hundreds
may be met with. This is not, however, the case with birds that
die quickly of acute disease.
Goniodes tetraonis is usually found on the smaller feathers,
crawling about halfway between their insertion and the tip of
their vanes. When disturbed they hurry away into the brush-
wood of the small feathers, like small] deer seeking cover, and
they are by no means so easy to catch as one would at first think.
They eat the finer barbules of the feathers, which, accumulating
in the crop, gives the dark curved marking in thew rather trans-
parent bodies. On this meagre and arid diet they seem to flourish,
actively produce young, and pass through several ecdyses.
The naked-eye colour of Goniodes is a yellowish brown. Under
the microscope the body appears rather transparent, but wherever
there is chitin this is of a yellowish to chestnut-brown colour
according to the thickness. The crop, which is full of minute frag-
ments of the finest barbules of the feathers, presents a blackish
sac-like appearance, running obliquely across the middle line of the
abdomen ; asomewhat parallel but much smaller black tube repre-
sents possibly the rectum (Pls. XL.& XLI.figs.12&16). In afew
cases the cesophagus and crop presented a red appearance, this
being probably due to hzemoglobin from the blood of the grouse.
* “Insecta Epizoa,’ Leipzig, 1874.
t Leiden, 1880.
af Zeitsch. f. ges. Naturwiss. xxviii. 1866, p. 387.
§ Chapman & Hall, London, 1877. :
312 DR. A. E. SHIPLEY ON [ Mar. 16,
The body is, on the whole, flattened—especially is this the case
with the head and abdomen. The thorax, as Snodgrass* points
out in Menopon persignatum, appears to be triangular in cross-
section.
The Head.—The head is shaped somewhat like the semicircular
knives used for cutting cheese. The head of the female is some-
what broader and shorter than that of the male, and is produced
at the posterior-lateral region into a much more prominent angle.
In both male and female the angle bears a spine and a long hair.
The anterior rim of the head is bounded by a thick rim of chitin,
beneath which is a layer of granular protoplasm with a few nuclei,
the hypodermis. At intervals the chitin is pierced by narrow
channels, into which the hypodermis extends, and the chitin bears
at the outer end of each of these channels a short hair (Pl. XXXVI.
fig. 4). From the inner surface of this rounded anterior edge
of the head a number of muscle-fibres pass radially inwards to a
structure which has been called the upper lip, and which will be
described later under this name. Just in front of the recess
from which the antenne arise, the anterior thickened chitinous
rim curves to an end, being bent in and then out to form a short
apodeme of which the inner end acts as the basis of articulation of
the anterior limb of the stout mandibles(Pls. XXX VI.& XXX VII.
figs. 4 & 6). The socket of the antenne is also provided with a
thickened chitinous skeleton, and across the base of the head,
separating it from the prothorax, is a thickened plate which
presents in profile the appearance of a bow; the rest of the head
is enclosed in thin yellow chitin. The appendages of the head
will be described later, together with those of the body.
There is no neck, but the first segment of the prothorax is
only about one-half the width of the head. The mesonotum
is fused with the metanotum and the thorax appears to have but
two segments. There is, again, no waist or constriction between
the thorax and the abdomen, but the segments from the first
thoracic to the second or third abdominal gradually and uniformly
widen, and then as uniformly diminish in width until the last.
The separation of the first thoracic segment from the second is
marked by a stout chitinous rim both dorsally and ventrally—
this is, however, only found in this region; the rest of the
segments are soft and not chitinized in the tergal and sternal
regions, but the pleura are protected by well-marked chitinous
shields, which, however, extend but a very short way dorsally and
ventrally (Pl. XXXYV. figs. 1,2 & 3). In Virmus, however, the
lateral plates are more extended.
According to Sharp? the Mallophaga have from 8 to 9
abdominal segments, and according to Railliet £ the family in
which he places Goniodes has 9; but he remarks that the last
* Pap. Calif. Ac. vi. 1899, p. 146.
+ Cambridge Natural History, vol. v. Insects, i. London, 1895.
t ‘Traité de Zoologie Médicale et Agricole.’ 2nd Edn., Paris, 1895.
1909.] ECTOPARASITES OF THE RED GROUSE, ilies
two are sometimes completely fused, so that we only find 8 visible
segments.
There are certainly only 8 visible in Goniodes, although
Nirmus has 9 complete segments. The last visible segment in
the female is a slightly bilobed plate bearing no hairs; the anus
opens just below it. In the male the plate is not bilobed; it is
stouter and bears a number of backwardly projecting hairs.
Each segment, except the last in the female, bears a number of
hairs; the arrangement of these is shown in the figures 1, 2, & 3,
Plate XXXV.
The appendages are as follows :—
I. The eyes.—Each eye is formed of a little aggregation of
pigmented cells, the whole somewhat cup-shaped, and of an
almost spherical transparent thickening of the cuticle, the lens.
The eyes are situated close behind the thickened cavity from which
the antenne arise (Pls. XXXV. & XXXVI. figs. 1, 2, 3, & 4).
JJ. The antennze.—These arise from 2 deep hollow the chi-
tinized walls of which are much thickened. The cavity practically
conceals the proximal joint, which is broader than long; the
second joint is the longest and is almost twice as long as its
broadest part; the third, fourth, and fifth segments uniformly
diminish in size, and the fifth or last bears at its end a number
of bristles. The male is readily distinguished from the female
by the fact that the third joint is produced into an inwardly
directed process very like a thumb, and this gives the antennz a
biramous appearance (Pls. XXXV. & XXXVI. figs. 1 & 4).
The next three pairs of appendages are modified as mouth-
parts, and in describing them we propose to mention certain
median structures also connected with the mouth.
The most remarkable feature of the under surface of the head
of a Goniodes is a white cushiony area with the outline of a
stout sausage, sometimes described as the “upper lip” or
“labrum.” It is bounded anteriorly by a ridge of chitin which
takes its origin on each side from the inner face of the strong
apodeme already mentioned. This cushion is covered with a
multitude of rugosities, giving it the appearance of the skin of
a dog-fish. There is always a more or less well-marked crease
or groove across the long axis of the cushion, and the part
posterior to the crease is supported by two longitudinal bars of
chitin just as the double banners Temperance reformers carry in
their processions are supported by the poles (Pl. XXXVI. fig. 5).
If one be watching the living Goniodes lying on its back on a
slide, this cushion will be seen from time to time to swell up and
scrape along the under surface of the cover-slip. Then it subsides
again, possibly being pulled back by the numerous muscle-fibres
which pass back from the anterior end of the head, and which
appear to be inserted into the inner surface of the cushion.
Along the posterior edge of the cushion is a small mobile mem-
brane or lip which bears a moustache of eight hairs, shorter in
Proc. Zoou. Soc.—1909, No. XXII. 21
314 DR. A. E, SHIPLEY ON [ Mar. 16,
the centre, but increasing in length as one passes outward
(Pl. XXXVI. fig. 5). This lip is frequently drawn down over
the tips of the mandibles.
The only function one can suggest for the upper lip is that it
acts as a scraping organ and it may be of use if the animal ever
eats the epidermis of its host.
III. The mandibles.—These are by far the most powerful of
the mouth-parts and are very strongly chitinized. The right
and left mandibles are not exact images of one another, as the
tip of one always closes outside the tip of the other, and thus
there is a slight differentiation at the apex, which is so strongly
chitinized as to be almost black. Hach mandible is somewhat
triangular in shape, the apex forming the tip. The articulation
is very complex (Pl. XX XVII. fig. 6) and is mainly with the
inner end of the powerful apodeme which runs in just in front of
the base of the mandibles. A very powerful muscle runs into
the external posterior angle of each mandible, the so-called
condyle, and serves to bring it into biting contact with the other.
The sharp shearing-edge of the mandible is admirably adapted
for cutting off the barbules of the feathers which form the food
of the biting-lice.
IV. The first pair of maxille.—These are very difficult to see
in the living animal and are best observed when in movement.
We agree with Grosse* in describing them as lobes without any
traces of palps. They are rounded and bear certain sete on them.
When in motion they are shot up and pulled down between the
mandibles and the labium or fused second maxillze; sometimes
both are moved forward at once, sometimes they move alter-
nately (Pl. XX XVII. fig. 7).
V. The second pair of maxillee—These have fused together and
form a labium of a very simple kind. There is a median plate or
mentum in which we found no transverse furrow. This plate
bears anteriorly a pair of one-jointed processes ending in a few
short stiff bristles. These are called by Grosse the paraglosse,
but, as there are a pair of minute one-jointed processes internal
to these, it may be that they represent the palps. Whichever
they are, they are very mobile and are constantly being divaricated
into a position at right angles to the normal and then suddenly
brought back again. They are obviously of use in bringing food
to the mouth. The more median processes as well as the palps bear
hairs. They are figured on Pls. XXXVI. & XXXVII. figs. 5 & 7.
A median structure which we think may represent the hypo-
pharynx is the lyriform organ, or the ‘“ cesophageal sclerite” of
Kellogg. his median piece is strongly chitinized, deep brown
in colour, and consequently conspicuous ; it seems to lie about in
the same level with the first maxille, except when they are
* Zeitschr. wiss. Zool. xlii. 1885, p. 587.
1909. | ECTOPARASITES OF THE RED GROUSE. 315
protruded, when it lies behind them. A muscle on each side of
the esophagus runs from the anterior angle of the sclerite to the
dorsal side of the head and brings about the movement of the
organ. Kellogg has described in certain species a pair of oval glands
which le ventral to the sclerite and the ducts of which unite and
open by a common duct into the median groove of the thickening.
These glands are very conspicuous in Goniodes, and are shown in
several of our figures. Their function is unknown. Their ducts
are cross-barred like a trachea. The whole sclerite is conspicuous
and shines through as a somewhat V- or U-shaped dark area,
visible from above. As Kellogg points out, a similar apparatus
exists in the Psocide (Pls. XXXV., XXXVI., & XXX VIL. figs. 3,
5, & 7). Two salivary glands on each side of the cesophagus have
been described in many genera of Mallophaga. The ducts of all
four unite and open into the pharynx by a common duct.
The second maxille are so minute and feeble that we found it
impossible to dissect them out even from macerated specimens.
VI. The prothoracic legs.—These pair of appendages are turned
forward and their ends normally lhe underneath the mouth. They
doubtless take some part in feeding. Their inner ends are ap-
proximated, so that the sternum here is but little more than
a line. Snodgrass* records that the prothoracic legs do not
move synchronously with either of the others or with each other.
One often moves backward as the other moves forward, and he
holds that they serve to guide the body. He thinks they serve
to pull the body up the feather, the prothoracic legs pulling
whilst the other legs push, like a man climbing a rope.
VII. The mesothoracic legs.—These are larger than the pre-
ceding and are directed backwards ; their bases are further apart.
The details are shown in Pl. XX XIX. fig. 10. All the legs end
in claws and bear a well-marked pulvillus.
VIII. The metathoracic legs.—These are still larger and, like
the preceding, are directed backwards; the sternum between their
bases is rather wider. The right and left mesothoracic legs move
forward simultaneously and backward simultaneously, and so do
the right and left metathoracic legs; but when the mesothoracic
legs move forward the metathoracic legs move backwards, and
vice versa.
In their general structure there is little beyond size in which
the legs differ. Each consists of a coxa firmly applied to the
ventral surface of the thorax; it is a broad, short piece, wide
distally. The second article is a small trochanter which joins the
hinder end of the wide coxa and seems to be almost part of the
femur, but there is a marked thinning of the cuticle between it
and the femur and a clear joint.
The femur with the trochanter and the next article or tibia
* Toc. cit. p. 152.
21
316 DR. A. E, SHIPLEY ON | Mar. 16,
are of about equal length, but the tibia is not so stout; distally
it bears a pair of stout bristles, hardly movable, against which the
tarsal claws work. There are other bristles on the tibia, and
numerous hairs on all the articles. The tibia bears a single-
jointed tarsus which carries a pair of very mobile claws. These
claws are constantly being depressed, usually one at a time, and
rub against the tibial bristles. The tarsus also carries numerous
knobs, and between the base of the claws a pulvillus may be seen ;
this in some cases is retracted.
The female has no external organs of reproduction, but on the
seventh segment of the male there are situated ventrally a couple
of complicated gonapophyses which presumably are modified
abdominal’ appendages. These are figured on Pl. XXXVIII.
fig. 9, which gives an adequate idea of their complexity.
Respiratory Systeno.
The tracheal system of Goniodes opens on the exterior by seven
pairs of stigmata. There may possibly have been more, but we
could not detect them. The most anterior is the largest; it is
situated close behind the first pair of legs and is very difficult
to see. Snodgrass * has described one in a similar position in
Menopon titan. From it a trachea passes inwards and gives off
a twig to the second leg. Further on it divides, one stout branch
running anteriorly into the head, where it divides into two, each
splitting up into innumerable small branchlets supplying the
organs in the head. From the main trachea the second branch
passes backward, giving off a twig to the third pair of legs, and
then runs backward through the abdomen as a main longitudinal
trunk (Pl. XX XIX. fig. 11).
The abdominal stigmata are twelve in number, there being a
pair on the second to the seventh segment, both included. They
lie on little eminences like a tee in a teeing-ground, situated
about one-sixth of the body breadth from the edge, and from
each is given off a short trachea which soon splits into two
branches. Of these the posterior splits up into innumerable fine
twigs, which supply the various organs of the segment, and the
anterior runs almost straight into the longitudinal trunk, thus
placing the system connected with one stigma in communication
with all the others on the same sides of the body. By this means,
if one stigma be blocked the organs it supplies are not deprived
of air, but receive it from another system. The smaller tubes
on each side pass across the middle line and seem to place the
right and left systems in communication. In Menopon titan,
according to Snodgrass, the right and left systems communicate
by means of a large transverse trunk in the fourth abdominal
segment.
The spiral thickenings are well marked.
* Loe. cit.
1909.] ECTOPARASITES OF THE RED GROUSE. 317
The Alimentary Canal.
Grosse has described just within the mouth a dorsal and a
ventral piece of a “schlundskelet.” Unless the lyriform organ, or
“ @sophageal sclerite,” represents the ventral] piece, this structure
is not evident except in sections.
The esophagus is a simple tube with muscular walls which
traverses the posterior part of the head and the thorax (Pl. XL.
fig.12). Soon after it reaches the abdomen it gives off a blind
pouch or crop, which is always choked with feathers and forms
the conspicuous black patch which shines through the wall of the
abdomen. The walls are very muscular, both longitudinal and
circular muscle-fibres being conspicuous. It usually lies near
the middle line, but slightly obliquely, and pointing posteriorly
to the right. Behind the point where the crop is given off the
stomach or chylific ventricle passes backward, lying to the right
of the crop. At the posterior end of this the four Malpighian
tubules arise, and then there follows a short intestine in which
usually masses of undigested feather-fragments are to be seen.
The intestine is short and ends in a ring of six almost spherical
(? glandular) bodies (Pl. XL. fig. 12). Each of these seems to
consist of a single gigantic cell, and the whole is very richly
supplied with tracheez. These bodies closely resemble similar
structures found in the rectum of many Diptera, ¢. g., the blow-
fly and the mosquito. Behind them there is a short rectum,
which ends in an anus situated beneath the terminal plate.
Numerous muscles run from the body-walls of the last two
segments to be inserted into the rectum and doubtless act as
divaricators.
The food consists of feather-barbules; a sample of it taken
from the crop is shown in Pl. XL. fig. 13.
The Exeretory System.
This system consists of (a) the Malpighian tubules, and
(6) the Fat-body, in which nitrogenous waste matter is often
stored away. The Malpighian vessels are four in number; they
arise at the anterior end of the intestine, and near the base each
swells into an oval vesicle (Pl. XL. fig. 12). The tubules are
long, as long almost as the body, and are coiled away amongst the
viscera.
The fat-body is very definitely arranged, there being paired
pouches of it at the sides of each segment (Pl. XLI. fig. 16). In
the cavity of these pouches are five collections of oval structures,
which may be the five pairs of ovarian tubules, showing the ova,
but somewhat similar structures occur, in equal numbers, in the
taale abdomen.
318 DR, A. E. SHIPLEY ON (Mar. 16,
Nervous System.
We have not made a detailed examination of the nervous
system, but may remark that it consists of a brain and a large
infra-cesophageal ganglion in the head and of three ganglia in
the thoracic seoments, The last of these is the largest, and
it supplies nerves to the organs of the abdomen.
Circulatory System.
This, again; we have not examined, but Wedl* and Kramer?
have seen and described the hearts of several species. They
seem to conform to the usual insect type, but the number of
chambers is small, Wedl says only one in Menopon pallidum,
situated in the last but one abdominal segment.
Reproductive Organs.
We have not investigated this system of organs in any detail,
but it may be mentioned that in the Ischnocera, the subdivision
of the Mallophaga to which Goniodes and Nirmus belong, there
are four testes, the two on each side being united by a common
vas deferens, which leads into a vesicula seminalis, which, though
bilobed, is usually unpaired; from this an ejaculatory duct leads
to a retractile penis. The extreme complexity of the external
male organs is shown in fig. 9, Pl. XXXVIIT. Morphologically
there is an invagination of the body-wall of the last abdominal
segment to form the genital cavity, and the various plates and
bars which are seen in the drawing are chitinous thickenings in
the walls of the invagination. In the centre of the genital
cavity lies the penis, which is strengthened by chitinous rods and
bars, and is capable of being protruded and retracted by a com-
plicated system of muscles. In the male the anus has been
involved in the invagination and comes to open dorsally into the
genital cavity. This is not the case in the female, where the
invagination is not close to the posterior end, but is formed by
an invagination of the eighth abdominal seoment. The vagina
opens anteriorly and dorsally into this chamber, and passes into
a long coiled oviduct which splits into two collecting- ducts, and
these terminate in five ovaries on each side of the “body. The
ovaries dwindle out anteriorly, and their thread-like forward
ends unite into a common termination.
An excellent comparative account of the reproductive organs
of the ern is given in Snodgrass’s already-mentioned paper,
and Gross £ has written an necount of the histology of the ovary,
which he finds strikingly like that of the Pediculidee.
* SB. Ak. Wien, xvii.
+ Zeitschr. wiss. Zool. xix. 1869, p. 452.
t+ Zool. Jahrb. Anat. xxii. 1905, p. 347.
1909. | ECTOPARASITES OF THE RED GROUSE. 319
The Eggs.
The eggs are very beautiful objects ; in badly infested grouse
they may be numerous, but as a rule they were none too easy to
find. Usually they occur in small groups attached to the base of
the after-plume and between it and the shaft of the plume.
The specimen figured was on one of the feathers from the
flank.
The eggs are elongated, some three to four times as long as they
are broad. ‘They are fixed by some adhesive secretion at the end
corresponding to the posterior end of the contained embryo. At
the other end is a well-marked cap or operculum which always
points to the free end of the feather. The beauty of the re-
ticulated egg-case is shown best in the genus WMenopon, and we
figure one, which we take to be the egg of Menopon pallescens
Nitzsch, found on the feathers of a partridge. Under the pressure
of a cover-slip the egg-case gradually ruptured along a circular
line below the well-marked thickened edge or rim of the
operculum. ‘The contained egg then began to emerge, carrying
the operculum as a sort of cap, the resemblance to which was
emphasised by the long process which stands out like a feather
borne on the apex. The eggs of Goniodes show the reticulations
less well, but they are well marked on the operculum, which bears
a long tapering filament, longer than the egg itself. They also
occur just below the opercular rim, but fade away towards the
fixed end. The general appearance of the eggs in the after-plume
is shown in P]. XL. fig. 15. They were found on the 27th July,
1908, and they seem to be laid throughout the summer.
There is no metamorphosis, the young leaving the egg-shell as
a miniature of their parents.
Il.—Wyruus CAMERATUS Nitzsch.
This insect seems to have been first named by Nitzsch * in the
year 1818, but with no description. Indeed, the animal is men-
tioned under the subgenus WVirmus, but is called Philopterus
cameratus. It is figured and described, and a bibliography is
given, in Denny’s ‘ Monographia Anoplurorum Britanniz’ 7 under
the name of Mirmus cameratus. Denny found it on the Red
Grouse, the Black Grouse, ‘“‘ and I expect also on the Ptarmigan.”
Grube describes it in Middendorft’s ‘ Siberian Travels’ as existing
on Lagopus albus and L. alpinus, thus confirming Denny’s
surmise.
It is mentioned in Giebel’s articlet on the Halle Bird-lice,
and described and figured in his great monograph ‘ Insecta
* Germar’s ‘ Magazin der Entomologie,’ Halle, iii. 1818, p. 291.
+ London, 1842, p. 112.
t Zeitschr. ges. Naturwiss. xxviil. 1866, p. 370.
320 DR. A. E. SHIPLEY ON [Mar. 16,
Epizoa.’ Piaget, in his ‘ Les Pédiculines,’ states his conviction,
that WV. cameratus is specifically identical with the V. quadrulatus
of Nitzsch, from Tetrao wrogallus, the Capercaillie. Kellogg in his
Mallophaga (‘ Genera Insectorum ’) does not mention WV. cameratus
though he records V. quadrulatus from 7. urogallus, T. tetrix,
and Lophophorus impeyanus.
NVirmus is a more slender animal than Goniodes, and appears
to be longer. It is rarer than the latter, though in the great
majority of cases the two are found together. Most of what has
been said above about Gloniodes applies also to Virmus, as their
habits are very similar, except that Virmus lives more on the
skin and upon the base of the rachis of the feather than does
Goniodes. It also seems to frequent the feathers under the wing,
where Goniodes is seldom seen. Both species seem to wander all
over the body ; and though they seem rather more common upon
the head, neck, and back, the old view that these biting-lice
oceur chiefly or exclusively on those parts of the body inaccessible
to the beak was not borne out by our investigations (Pl. XLI.
figs. 17 & 18).
The variation in size and in colour is very considerable. Dead
specimens are not infrequently found, and these may be in some
cases mistaken for cast skins. An average length is 3 mm., and
an average width of the abdomen is 15mm. The abdomen is
the widest part. In no case did we find either Goniodes or
WVirmus in the crop of the grouse, though, as we have just stated,
they are fully exposed to being snapped up by the bird’s beak if
the bird cared to notice them. It is not known exactly how
clean birds get infected: probably the Mallophaga simply crawl
from one bird to another when the latter are contiguous, and the
young birds are infected on the nest. There is evidence, however,
that in some cases, probably rare ones, they cling to the grouse-
fly and are by it transported to a new host.
In the summer of 1907 Mr. Fryer found some Mallophaga
eggs. These were for the most part empty, but from one or two
full ones he has succeeded in hatching out specimens of Virmus
cameratus. ‘The eggs are white, and transparent when empty,
just visible to the naked eye, 0-6 mm. in length, and about four
times as long as they are broad. Each egg-case is beautifully
reticulated, the areas between the reticulations being six-sided.
At one end the egg has a cap which is pushed off when the young
emerges. The eggs are laid between the barbules of the vanes or
near the bases of the filo-plumes, and adhere to their supports by
means of some sticky excretion (Pl. XLII. fig. 20).
The eggs appear to be laid throughout the summer; the first
time we found them (some of them were empty) was on 2nd July,
1907, and we found others later in the season.
There is no metamorphosis ; the young emerge from the egg-
case as small miniatures of their parents. They seem to cast
their skin several times, but the exact number of ecdyses is not
known.
1909. | ECTOPARASITES OF THE RED GROUSE. 321
B. Diprera.—F lies.
a.) Fam. Hippoboscide.
TIl.—OrayrHomyra LAGoPopIs Sharp.
Till recently it had been thought that the grouse-fly was the
same species as the common bird-fly, Ornithomyia avicularia L. ;
but recently Mr. D. Sharp* has pointed out that it is a distinct
species, which he has described, as follows, under the name of
O. lagopodis:—It is “smaller than O. avicularia, and distinguished
by its peculiar lurid blackish colour, without any trace of green
even on its feet or legs; the rostrum is black, and the hairs of
the body and appendages are shorter than in the better-known
form; on each side of the thoracic pleuron, between the front
and middle legs, there is a very large dark patch extending as
far towards the middle as the base of the front coxa, and divided
into two parts by an oblique pallid line. The head is considerably
smaller and narrower than that of O. avicularia, and has beneath
a verv large area of smoky colour on each side. Mr. Collin has
pointed out that the segments, or abscissze, of the costa afford a
good character ; the relative lengths of the outer two being in
O. lagopodis as 9-8, and in O. avicularia about 12 or 123-8.
The bristles on the scutellum are usually more numerous, as well
as larger, in O. avicularia.” Recently a second species, O. frin-
gillina Bezzi, has been separated off from the O. avicularia, so that
we now have three species of Ornithomyia in this country, and
probably more will be added as the group is further studied.
Mr. Sharp thinks that the same species frequents the willow-
grouse, L. albus, of Scandinavia.
The head and mouth-parts of this fly are very interesting.
A ventral view shows, between the eyes, the short antennz
apparently of two joints, ending in four hairs, of which one is
far longer than the others; other symmetrically arranged hairs
are shown in Pl. XLIV. fig. 22. In the middle line is the
proboscis; this consists of two lateral, movable, palp-like struc-
tures, each bearing hairs and terminating in a stout bristle.
These structures are presumably the maxillary palps. Then there
is a median very mobile structure, which is the sucking-tube ;
this moves in all planes, and may be protruded or withdrawn.
Tits mouth shows a somewhat plicated orifice, and behind it
undoubtedly endsin a sucking pharynx. This median structure
is probably homologous with the second maxille or the labium.
The feet of the grouse-fly are large but very beautiful. In
Pl. XLIV. fig. 23, we show the outer surface of one of the feet,
and the same figure shows part of the femur, the tibia, and the
five joints of the tarsus, the fifth being by far the largest.
* Ent. Monthly Mag. II. ser. xviii. 1907, p. 58.
322 DR. A. E. SHIPLEY ON [ Mar. 16,
Fig. 24 shows the inner surface of the same, and on Pl. XLIII.
will be found a coloured sketch of a foot seen obliquely. From
these drawings, which have been prepared for me by Mr. Edwin
Wilson of Cambridge, it will be seen that the large paired
claws are double, and that whereas the distal limb of each claw
is slender and very sharply pointed, the proximal limb is much
stouter and ends bluntly. Between the claws is a median,
feathered, process with hairs or bristles, and at the base of each
double claw is a pulvillus covered with minute hairs. As fig. 24
shows, there are other processes for which I have no name.
The arrangement of the hairs is faithfully given in the drawings.
We do not know the exact relations of the grouse-fly to the
grouse. It is believed to suck its blood, and it will certainly bite
human beings. For a time it seems to burrow amongst the
feathers of the bird, and anyone handling grouse during the
summer is likely to disturb a fly or two. They come buzzing out
and are apt to crawl up one’s sleeve by aid of the pair of great
hooked claws on their feet. Altogether they have a sinister
aspect, and to people who do not like flies they are very repellent.
They occur freely in larders where freshly-killed grouse have
been placed, and after a short time they leave their dead host and
accumulate upon the windows.
The earliest month we have found the grouse-fly is in June,
towards the end. The latest we have found up till the present
time is September. Mrs. Duff Dunbar has taken them as late as
October. Perhaps they are most plentiful in August.
The females seem to be commoner than the males, or, it may
be, in August are more readily taken. Like other members of
the Hippoboscidee, which includes the horse-fly, forest-fly, and
sheep-tick, the grouse-fly does not lay eggs, but the ovaries pro-
duce one large ovum at a time, and this passes into a dilated
oviduct which acts as a uterus, and here the egg develops. After
attaining a certain stage of development, the larva surrounds
itself with a pupa-skin and is extruded. The chitin covering the
larva hardens and blackens with exposure to the air, and forms
the so-called pupa-case ; in fact, one may almost say the young
are hatched as pup. At no time is the larva exposed, though
there is a larval stage free in the uterus wrapped first in the
egg-sheil and then in the pupa-case.
When first deposited the pupz are light in colour and the case
has not hardened. Those dissected out from a fly are shorter
and more squat than the mature pupe found on the ground
(compare a and 6 in fig. 25, Pl. XLIV.), and the symmetrical
ridges and elevations are much less well marked (Pl. XLIYV.
fig. 25,d &c). The mature pupe are shown from above, from the
side, and from the micropylar end in Pl. XLIV. fig. 25, ¢, e, & f,
highly magnified. Between them they show well the six elevated
and cross-barred ridges which radiate from the micropyle to the
angles of the hexagonal micropylar area.
The pupz were found during August and September. They
1909. | ECTOPARASITES OF THE RED GROUSE. 323
appear to be deposited amongst the feathers and are easily
detached from them. The few we have found either dropped on
some paper over which we were handling some birds, or lay loose
at the bottom of the cardboard boxes in which grouse travel.
Probably they take some eight or nine months before they give
rise to the imagos, and the latter very likely disappear altogether
from about October till June. Further research is needed tc
throw light on these questions.
Three specimens of O. lagopodis, all of them taken from one
grouse, were themselves markedly infested with an ectoparasite,
a species of mite. Here I refrain from quoting Dean Swift.
The mite belongs to the genus Canestrinia, as my friend Mr. C.
Warburton has kindly told me, and is probably a new species.
The subfamily Canestrinine are all parasitic upon insects, and
are regarded as harmless. Our specimens existed in considerable
numbers, clustered round the hinder end of the fly’s abdomen on
the ventral surface, with their proboscides plunged into its body.
Many were laying eggs, and many cast-off cuticles were lying
around. Eggs from which the larve had escaped presented a
spindle-shaped outline; others contained ova in various stages of
differentiation ; others fully formed larvee.
We have in no single case found a grouse-fly in the crop of a
grouse, nor have we yet found any cestode larve or cysts in the
bodies of the flies which we have cut into sections or dissected.
Gi.) Fam. Scatophagide =Scatomyzide.
LV .—SCATOPHAGA STERCORARIA lL.
This fly cannot be looked upon as an ectoparasite of the grouse,
but it lays its eggs in grouse-droppings, and its maggots live on
and in these dejecta. The maggots must therefore constantly be
in close contact with and possibly eating the ova of the tape-
worms which exist in such vast numbers in the grouse-droppings ;
and here we thought it was a profitable object to investigate for
the cysticercus or second stage of the cestode. It should be
mentioned that the droppings consist of two parts: (1) the dejecta
from the intestine strictly speaking, and (2) the more fluid dejecta
from the ceca. The latter pass last and lie like a cap upon the
former. The fly-maggots are only found in numbers in the
“cecal” part of the dropping. Mr. Fryer first found them
commonly at Fort Augustus in April. In June they were not so
common, owing perhaps to the rain which washed the cecal part of
the droppings away. We examined a Jarge number of the larve
both by squashing them and cutting them into sections, but we
found no trace of infection; in fact, here, in this most likely
place, we again drew a blank. No specimen of S. stercoraria of
of its larve has been found in the crop. This fly, which, as
324 DR. A. E. SHIPLEY ON [ Mar. 16,
stated above, we believe to be S. stercoraria, may eventually turn
out to be a local variety.
The larva of the fly has the usual maggot-like shape, tapering
from behind forward towards the mouth. Counting what appears
to be the cephalic segment—but which in reality probably repre-
sents more than one segment, and which is thus conveniently
called the ‘ pseudo-cephalon ” *—as one segment, there are thirteen
in all, the usual number for Dipterous larvee (Pl. XLV. fig. 26).
The cuticle is thin, the maggots are white. ‘They bear numerous
small spines which are especially conspicuous in a ring around
the anterior end of each segment. These rings emphasize the
segmentation of the larva.
The ‘pseudo-cephalon” is pointed, and varies in outline
according to how much of the mouth and its sclerites are pro-
truded or not. These sclerites are the most conspicuous structures
in the larve; jet-black, they stand out against the white tissues
of the maggot. There is a pair of hooks which apparently
correspond with the single median mandibular sclerite of Musca
domestica. At the base of each of these is a dentate sclerite,
and behind the mandibular sclerite articulates with the hypo-
stomal sclerites. These latter are irregular longitudinal bars
connected by a slight transverse plate on the ventral side. In
M. domestica the salivary glands open into the pharynx just
in front of this transverse piece. Posteriorly the hypostomal
sclerites are very closely articulated, or perhaps even fused with
the large lateral pharyngeal sclerite. This consists of a ventral
plate, continuous with two lateral plates which are deeply notched,
and in the house-fly the nerves and tracheze which supply the
pharynx enter through this notch. The two lateral plates are
united anteriorly by a dorsal cross-piece. The whole of these
sclerites are being continually pushed forward and retracted by
a complicated series of muscles which have been carefully described
in the case of MW. domestica by Dr. C. Gordon Hewitt (Pl. XLV.
fig. 27).
The mouth is bordered by tumid lips, above which the hook-
like mandibular sclerites are pulled and pushed in and out.
Dorsal to these again are two elevations which each bear two
sensory papille; these correspond with the sensory tubercles of
AW. domestica, though the latter are not borne on an elevation.
The anus opens on the truncated thirteenth segment, well
forward on the ventral surface ; around it, symmetrically placed,
are four anal papille, which assist in the movements of the
maggot (Pl. XLVI. figs. 31 & 32).
The tracheal system opens on the flattened posterior end, about
the centre. Ventral to the stigmata there are two or three
pairs of low papille. Hach stigma leads into a trachea, which
almost immediately splits into a visceral branch (Pl. XLV. fig. 26)
which bends down into the viscera and extends a little way in
* Henneguy, L. F., ‘Les Insectes,’ Paris, 1904.
+ C. Gordon Hewitt, Quart. J. Mic. Sci. lii. 1908, p. 495.
1909. ] ECTOPARASITES OF THE RED GROUSE. 325
front of the middle of the body. Before splitting in I. domestica
the right and left main trunks are put into communication by a
transverse trunk ; if this exists in S. stercoraria it escaped our
notice. ‘The lateral trunks give off in each segment a dorsal and
ventral twig. Anteriorly, in what appears to be the third segment
(it is described as the fourth in the house-fly), there is a transverse
commissure by means of which the right and left trunks are put
into communication. In front two small twigs are given off from
this transverse commissure which run to the pharynx. The main
trunk is continued forward, and at the second segment (the third
in the house-fly) it ends in a process like a little rake (Pl. XLV.
figs. 26 & 27). This is due to the splitting up of the trachea into
eight or nine little twigs, all in the same plane and all ending in
a knob. The whole is called the anterior spiracle, and can be
protruded from the body and retracted. Hewitt states that in
the house-fly each of these knobs opens to the surface by a very
minute pore.
C. SIPHONAPTERA.—F leas.
Ga.) Fam. Pulicide.
V .— CERATOPHYVELUS GALLINULZ Dale*.
Synonym: Ceratophyllus (Trichopsylla) newsteadi t Rothsch.
I am indebted to my friend Mr. N.C. Rothschild for identifying
this flea, which is here recorded for the first time from the grouse.
It is a well-known bird-flea, having been found in the nest of the
hawfinch, Coccothraustes vulgaris, in that of the dipper, Cinclus
aquaticus, in that of the blackbird, Z’wrdws merula, the moor-hen,
Gallinula chloropus L., and others. In the thousands of grouse
which have passed through our hands we have found but one or
two specimens of this flea, all in 1906, and we have never found
a single specimen in the crop. Hence, although the dog-flea,
Pulex serraticeps P. Gerv., is said to be the intermediate host of
the dog tape-worm, Dipylidiwm caninum, 1t seems hardly possible
that the C. gallinule could play any part in the life-history of
the grouse tape-worms. In identifying a flea almost every hair
tells, and as C. gallinule has not been accurately figured before,
I take this opportunity of figuring it in both sexes.
V1.—CERATOPHYLLUS GAREI Rothsch.
This second species of flea was found in a grouse in 1907; we
only took one or two specimens. It is recorded by Evans { from
the nest of the water-vole, of the lapwing, Vanellus vanellus, and
of the ring-dove, Columba palumbus. Rothschild $ has found it
* N.C. Rothschild, Ent. Monthly Mag. IT. ser. xiv. 1903, p. 145.
+ Ent. Rec. xin. 1901, p. 284.
Ann. Scott. Nat. Hist. 1906, p. 163.
f Ent. Monthly Mag. II. ser. xii. 1902, p. 225.
326 DR. A. E, SHIPLEY ON [ Mar. 16,
in the nest of a water-hen, Gallinula chloropus, and he records it
as having been taken from Mustela erminea, MW. vulgaris, Sorea
vulgaris, Microtus glareolus and M. amphibius, and trom hedge-
clippings.
ARACHNIDA.
AcARINA.—Mites and Ticks.
G.) Fam. Ixodidee.—tTicks.
VIL.—TLxopes ricinus (L.).
This is one of the commonest and one of the oldest-known
ticks of Europe. In the British Isles it usually occurs on
hunting-dogs and is sometimes cailed the “ dog-tick”; the adult
stage is especially frequent on sheep, goats, and oxen, less common
on horses, dogs, and men. Myr. William Evans*, of Morningside
Park, Edinburgh, tells me that he has not found this species on
dogs—in his district the “ dog-tick” being Jiodes heawagonus
Leach, var. inchoatus Neum. On the other hand, the larvee and
the nymphs are common enough on birds, lizards, and small
mammals—in fact, on animals which live among and brush
against grass or heather. It is only in the nymph and larva
state that we found these ticks on the grouse. On each of the
infested birds the specimens were fixed on the chin or around
the eyelids—in fact, in such positions as the grouse cannot reach
with its beak. In parts of Ross-shire, especially in certain woods,
these ticks swarm in enormous numbers, and the keepers assure
us that they kill large numbers of young blackgame. Hence there
is nothing remarkable in finding this species from time to time
on the grouse, where its presence must be regarded as accidental.
The larval stages emerge from the eggs and probably crawl on to
the heather, and thence on to the grouse or other animals which
come in contact with the vegetation. We have found both
larvee and nymphs amongst the feathers, but in small quantities
and on rare occasions. We have never found it in the crop, and
it can hardly play any part in infecting the bird with tape-
Worms.
Ixodes ricinus, or the “ castor-bean tick,” as it is called in
America, is common in many parts of the world. It is reported
from sheep, goats, cattle, horses, deer, dogs, cats, foxes, ferrets,
hedgehogs, hares, rabbits, bats, birds, and man. ‘This tick occurs
most frequently during the spring and early summer, but can be
found in lesser numbers up till September and October, possibly
later.
Severe epizootics amongst fowl of spirillosis and of another
obscure but very often fatal disease have been described by
* Ann. Scott. Nat. Hist. 1907, p. 35.
1909. } ECTOPARASITES OF THE RED GROUSE. 327
Balfour* in the Sudan. The spirochete, probably Spirocheta
gallinarum, which causes the first-named disease is transferred
from one fowl to another by a tick, Argas persicus. The second,
and as yet rather obscure, disease is recognised by the natives and
by them associated with the presence of the Argas. We have
found no traces of such disease in grouse, and the recorded
number of ticks taken in the grouse is, except locally, so small
that they can hardly play any part in grouse-disease,
(ii.) Fam. Tyrogly phide.
VITL.—Arevrosivs rarin% (de Geer).
Synonym: Tyroglyphus farinw Gerv.
Mr. C. Warburton has kindly identified for us a small mite
which was found in considerable numbers on several birds and at
varying times of the year. Whilst very common at Easter time,
they were less abundant in July. Alewrobius farinw, sometimes
known as the flour-mite, occurs in great numbers on all sorts
of organic material—grain, straw, hay, tobacco, flour, cheese,
dead bodies, ete., etc. At times workmen handling corn, cats,
horses, etc., have suffered much cutaneous irritation and eruption
from the attacks of this mite. There seems no doubt as to the
species of this mite, but the authorities on these animals express
surprise that they should occur so commonly on the grouse. Our
specimens, some of which were taken on freshly killed grouse,
contained some red substance in the stomach, probably some
blood from the bird. There seems at present little reason to
incriminate this mite as the carrier of the tape-worm cyst. They
were, however, found by Mr. Fryer on a large majority of birds
which were especially searched with the view of finding mites.
Gu.) Fam. Gamaside.
IX.—-GAMASUS COLEOPTRATORUM (L.).
We have also taken this common, fawn-coloured mite off the
feathers of a grouse. It is usually found on beetles, but winters
under stones, and it is said to soon die if removed from the beetle
or from under the stone where they hide, unless they are kept
moist. The beetles they favour are usually burrowers in the
damp ground or under cow-dung. ‘They probably passed on to
the grouse from under stones.
* Brit. Med. Journ., 9th November 1907, No. 2445, p. 1330.
328 DR. A. E. SHIPLEY ON [ Mar. 16,
GENERAL DISCUSSION ON THE RELATIONS OF ECTOPARASITES
TO THE ENDOPARASITES OF THE GROUSE.
We have in the alimentary canal three species of tape-worm,
two of the genus Davainea and one of the genus Hymenolepis.
We know that tape-worms, with perhaps the exception of one
species, pass through two distinct and different animals known as
hosts. In one animal it lives as an adult, in the other as a larva.
The larval host is always, sooner or later, eaten by the host of the
adult, and then the larval tape-worm or cyst grows into the adult
tape-worm. It was with the hope of discovering the second or
larval host of the grouse cestodes that we began a laborious
research on the insects and arachnids which infest the grouse.
Unfortunately, little or nothing is known about the life-history
of any species of either Davainea or Hymenolepis. ‘The larval or
cystic stages of the former have in some few cases been said to
occur in insects and in molluses; the larva of the latter is thought
to live in an insect or a myriapod, or perhaps even more likely
some ‘‘ water-flea ” or other fresh-water crustacean.
With regard to these possible second hosts. We have never
found a myriopod in the crop of a grouse, and so far we have not
found any crustacea—though it must not be forgotten that these
are probably so small as to escape notice. We have found one
species of slug in the crop of a Staffordshire grouse, which
Mr. W. E. Collinge has kindly identified for us as Arion empiri-
corum Feérussac, a species of slug which is common on the
Staffordshire grouse-moors. He tells me that the slug un-
doubtedly belongs to the genus Arion, and almost certainly to
Férussac’s species A. empiricorum, a name J. W. Taylor, in his
‘Monograph of Land and Fresh-water Mollusca of the British
Isles’ *, includes among the synonyms of Arion ater (.). The
well-known difficulty of identifying slugs which have been pre-
served and which have lost their colour accounts for the slight
doubt that exists. Arion empiricorwm is very voracious and
practically omnivorous; it will eat almost anything, especially
decaying animal and vegetable matter, fungi, paper, weak and
injured worms and slugs, and—what is interesting from the
point of view of the grouse tape-worms and round-worms—it
devours the dejecta of other animals. It prefers the shady places
in moors and fields, and emerges into the open only at dusk or
when the day is cloudy or overcast. The following parasites
which may give rise to adult forms in the grouse have been found
in A. empiricorwm :—
Trematopa (Flukes) :
(1) Cercariacum limacis Duj.7
(2) Cercaria trigonocerca Dies.¢
* Leeds, part xi. p. 167. + Dujardin, Hist. nat. des Helm. p. 472.
+ Leuckart, Paras. d. Menschen, 2nd edit. 11. p. 86.
1909. | ECTOPARASIVES OF THE RED GROUSE. 329
Cestopa (Tape-worms) :
(1) Cysticercus arionis v. Sieb.*
(2) Cysticercus tenice arionis v. Sieb.T
Nematopa (Round-worms) :
(1) Leptodera angiostoma Duj. =
(2) Leptodera appendiculata Schneider §.
(3) Nematodum limacis atra v. Sieb. ||
(4) Pelodytes hermaphroditus, Schneider 4.
We have cut one of these slugs into sections, and have sought
diligently through them for cysts of tape-worms, but have found
none. This absence of infection, combined with the great rarity
of the slug in the grouse’s crop, seems to show that A. empiricorum
is not the second or larval host of the grouse cestodes.
Dr. Wilson and Mr, Fryer “tow-netted” some of the moor
streams in April 1907, and found a certain number of the
nauplius larva, probably of Cyclops, and a certain but small
number of adult Cyclops. The numbers were, however, meagre,
and tow-nettings later in the summer yielded an even more
unsatisfactory “bag.” None of the crustacea when examined
microscopically showed any cysts, and as they were few in number
and quite cyst-less, it seems improbable that the source of the
tape-worm infection lies here.
Mr. D. J. Scourfield, who kindly looked through some of these
tow-nettings, tells me he found the following species of Ento-
mostraca :—
CLADOCERA.
Chydorus sphericus O. F. M., the most abundant form.
Alonella nana (Baird, Norman & Brady), frequent.
Alonella excisa (Fisch.), frequent.
Acantholeberis curvirostris (O. F. M.), a fair number, with
some cast ephippia.
COPEPODA.
Cyclops nanus Sars, a few.
Cyclops languidus Sars, a single specimen only seen.
Cyclops vernalis Fisch., again only one specimen was seen.
T subjoin three more lists of tow-netted freshwater Entomostraca:
from three different lochs. These were collected and identified
by Mr. Wm. Evans, who has kindly put them at my disposal, and
they clearly indicate the sort of surface fauna which may be
obtained from the lochs on the Scotch moors in early autumn.
Von Siebold, Zeitschr. wiss. Zoo]. 11. 1850, p. 202.
Krabbe, Nye Bidrag, p. 5; and Villot, Ann. Sci, nat. 6th ser. xy. 1883.
Schneider’s Monograph, p. 157.
Schneider’s Monograph, p. 159.
|| Schweiz. Zeitschr. Med. 1848.
4 Zeitschr. wiss. Zool. x. p. 176.
Proc. Zoou. Soc.—1909, No. XXII. LD)
rt
330 DR. A. E. SHIPLEY ON [ Mar. 16,
Lisr I.
From Loch Rusky, a moorland loch a few miles from Callander,
which was tow-netted on the 16th September, 1906.
CLADOCERA.
Simosa vitula (O. F. M.).
Eurycercus lamellatus (O. F. M.).
Alonopsis elongata G. O. Sars.
Alona affinis Leydig.
Chydorus sphericus (O. ¥. M.).
OSTRACODA.
Cyclocypris globosa (G. O. Sars).
Pionocypris vidua (O. F. M.).
Notodromus monacha (O. F. M.)
Candona candida (O. F. M.).
COPHPODA.
Moraria brevipes (G. O. Sars).
ces
Cyclops viridis (Jurine).
Cyclops annulicornis Koch.
Cyclops serrulatus Fischer.
List [1.
From Peat-pools on grouse-moors on Ben Ledi, in South-west
Perthshire, Sept. 1908.
OLADOCERA.
Chydorus sphericus (O. F. M.), very abundant.
OSTRACODA.
Herpetocypris tumefacta (B. & R.).
Cypridopsis villosa (Jux.).
Potamocypris fulva (Brady).
Candona candida (O. F. M.}.
COPEPODA.
Attheyella zschokkei (Schin.).
Atiheyella cuspidata (Schm.).
Cyclops vernalis Fisch.
List T11.
From Loch-a-Chroin, north of Callander.
CLADOCERA.
Bosmina longirostris (O. F. M.).
Acroperus harpe Baird.
Alonopsis elongata (G. O. Sars).
Alona quadrangularis (QO. F. M.).
Alonella excisa ( Fisch.)
Chydorus sphericus (O. F. M.).
1909. ] ECTOPARASITES OF THE RED GROUSE. 33]
OSTRACODA.
Cyclocypris serena (Koch).
COPEPODA.
Diaptomus gracilis (G. O. Sars).
Cyclops viridis (Jur.).
Cyclops serrulatus Fisch.
Also the common freshwater Amphipod, Gammarus pulex
(De Geer).
A complete list, so far as was known at that time, of the
Entomostraca of the Highlands and of the Lowlands could be
extracted from the very useful Synopses published by Scourfield
in the Journal of the Quekett Microscopical Club during the years.
1903 and 1904,
In none of the species examined have we yet succeeded in
finding any cysts.
We have thus with some degree of probability shut out as the
second or larval host of the tape-worms—at any rate for the
present—the ectoparasites of the grouse, the myriapoda and
the slugs or snails, and the fresh-water crustacea, and this on the
grounds (1) that on examination none of them reveals a cyst, and
(2) that these animals are either not eaten by the bird, or so rarely
eaten and in quantities so small as to render it highly improbable
that any of these invertebrates could account for the almost con-
stant presence of the cestodes in large numbers in the grouse.
Two rather striking facts seem to point to the normal insect
food of the grouse, which it picks up on the moor, as the more
probable source of the tape-worms. One is, that two of the
artificially reared grouse at Frimley, which died during the early
autumn of 1907, were carefully searched for tape-worms; but
neither Davainea nor Hymenolepis was found. ‘lhe second fact
is, that the young grouse often contain fully grown Davainea
before they are three weeks old. They must certainly have
swallowed the second host when very young, perhaps even the
day they were hatched, or the worm would not have had time to
erow. Hence our best chance of finding this second host is to
examine the crop-contents of the very young birds, and to do this
we must have a moor at our disposal, and leave to kill as many
young birds as we may want.
I have been assured over and over again by sportsmen and
gamekeepers that the grouse eats no insects, but this is far from
the truth.
Although the observations on the animal food of grouse are
still incomplete, enough has been done to show that it is fairly
abundant and very varied.*
From the crop of a single bird I have taken six larve of
* A fuller report on the insects found in the grouse-crop is given by Mr. J. C.
F¥. Fryer in the Interim Report of the Grouse Disease Inquiry, published in August
1908. The following two paragraphs relate to some observations of my own, made
in 1905 and 1906.
a7
B32 DR. A. E. SHIPLEY ON [ Mar. 16,
Tenthredinide (saw-flies), eight caterpillars of a Geometrid moth,
one caterpillar of a smaller moth, two small Tineid moths, a
number of immature Homopterous insects resembling the “ frog”
or “ cuckoo-spit,” a fly, possibly a Leptis, two specimens of the
family Aphid (plant-lice), one small spider, and the remains of
four specimens of the slug Arion empiricorum Fer. The gizzard
of the same grouse contained, in a more broken up condition and
consequently more difficult to identify, two or three dozen larvee
of saw-flies and moths, some young Homopterous insects, and the
pupee of two Muscid flies.
The segments of the grouse tape-worms containing the ripe
eggs pass away with its dejecta and get on the ground or on to
the heather and other plants, or ito water. The eggs of the two
species of Davainea are believed to develop into the cestode larva
inside the body of an insect or a land mollusc. They are ex-
cessively minute, and lying as they do in millions on the heather,
may be readily consumed by the leaf-eating caterpillars and other
insect larvee which live on the moors. Doubtless many are eaten
by the grouse themselves, but they are digested and come to
nothing. As we have said above, a tape-worm must have a
‘second or intermediate host, and its larval stage must be passed
inside an animal quite distinct from that which harbours the
adult worm. To get at and eat the eggs seems to me an easier
matter for caterpillars and other insect larvee or for slugs than it
is for the ectoparasites, which as a rule are not very likely to
come across the dejecta of their host. For this reason, in looking
for the larval tape-worm, we are now searching the insect larvee
and the slugs eaten so eagerly by the grouse. A common food of
grouse is the head of certain species of rush. Juncus articulatus
v. lamprocarpus, J. squarrosus, and J. effusus v. conglomeratus are
all frequently eaten. There is a very minute moth the larvee of
which live in curious, white, papery cases inserted into each twig of
the rush-head which they eat.. When the rush is in its turn eaten
by the grouse, the larvee of the moth pass into the alimentary
canal of the bird and are there digested. It has not been possible
to finally determine the species of the moth, but I think it is
Coleophora ceespititiella*, for this species frequents many species of
rush; whereas the C. glaucicolella, the other inland species, is
most partial to Juncus glaucus. The former is usually fully out
by the middle of June and lingers on till the middle of July ; the
last-named moth issues about the middle of July, and flies for
four weeks. The case is whitish, semi-transparent, and with
brown specks; it is found when the larva is no longer young, but
not at any very fixed time. At first its outer end is closed. The
larva often leaves the case, burrowing into the rush-head for food,
and at times fails to refind it. Before pupating, the outer or
anal end of the case is opened and the case strengthened by a
glandular excretion. These larvee should be searched for cysts.
* J. H. Wood, Ent. Mag. II. Ser. iii. (xxviii.) 1892.
1909. | ECTOPARASITES OF THE RED GROUSE. Boo
° EXPLANATION OF THE PLATES.
List oF ABBREVIATIONS.
a., anus. i | mi., micropyle.
ant., antenna. | m.s., mandibular sclerite.
a.p., chitinous rod supporting upper | m.t., Malpighian tubules.
movable lip. | Imax. & 1mx., 1st mavyilla.
a.s., anterior spiracle. | 2max. & 2mx., 2nd maxilla.
C., COXA. | mx.p., maxillary palp.
cer. In fig. 12, & c. in fig. 16, crop. ce., cesophagus.
dg. in fig. 27 & ds. in fig. 28, dental | Ov., Ovary.
sclerite. | p. & ped., pulvillus.
dg. & d.gl., ducts of oval glands. | prob., proboscis.
e@., eye. p-S., posterior spiracle.
emp., empodium. | ph.s., pharyngeal sclerite.
f., femur. | yx.eld., rectal glands.
f.b., fat-bodies. Sp., spines.
gl. & glds., oval gland. | St. im fig. 9, crop; in fies. 29 & 80,
h.l., “moustache” lip. | st., sensory papilla.
h.s., hypostomal sclerite. | sti., stigma or spiracle.
int., intestine. sto., stomach.
Ist lg., Ist leg. t.c., transverse commissiire.
lu., upper lip. | ta., tarsus.
lyr., lyriform organ. ti., tibia.
m., mouth. | tr., trochanter.
m.a., micropylar area. | iu.l., upper lip.
mb., rectum. | v.t., visceral trachea.
md., mandible.
PratE XXXV.
Fig. 1. Goniodes tetraonis Denny, male seen from above. The legs are shown on
the left side only. The forked character of the antennz, the upper lip.
and the lyriform organ are well shown in the head and the male genital
plates in the abdomen.
2. Goniodes tetraonis Denny, female seen from above. Note the different
shape of the head and of the outline of the body. The legs are not shown.
3. The same seen from below, showing the upper lip, the mandibles, the
Fig. 4.
5.
Fig. 6.
lyriform organ, and the legs.
Prate XXXVI.
Ventral view of head of male G. tetraonis, showing the forked antennz,
the mandibles, the pores in the chitin under the hairs, the muscles
running to the upper lip, the eyes, the lyriform organ, and the ducts of
the oval gland.
View of mouth-parts showing the “upper lip” w./., the moustache lip h.1.,
the mandibles id., the first maxille 1 maa, the second maxille 2 maw., the
ducts of paired glands d.g/. and the lyriform organ lyr., and the chitinous
bar supporting the upper lip a.p.; glds., the oval gland.
Pratt XXXVIT.
View of the ventral surface of right mandible showing the complex nature
of its articulation.
7. View of the 1st and 2nd maxillx and of the mouth, the mandibles having
Fig. 8.
)
Fig. 10
11
been removed. The glands with their paired ducts uniting in the middle.
line to open by a median duct into the pharynx, in the centre of the.
lyriform sclerite. mm. mouth.
Pratt XXXVIII.
View of the glands and the lyriform organ, showing the opening of the
duct.
Ventral view of the complex, male, external reproductive apparatus, formed
by various sclerites in the wall of the invaginated genital pouch. The
posterior angle of the crop st. is shown.
PLATE XX XIX.
. A ventral view of the right mesothoracic leg, showing the joints, claws,
and p. the pulvillus, ec. coxa, f. femur, ta. tarsus, fi. tibia, tv. trochanter.
. View of the tracheal system of a female, showing the stigmata, sfi.
DOA
ON ECLOPARASITES OF THE RED GROUSE. | Mar. 16,
Prate XI. °
Fig. 12. Alimentary canal of G. tetraonis dissected out: oe. cesophagus, ec”. crop,
Fig.
Fig.
Fig.
Fig.
13.
14.
15.
Ore
33.
b4.
sto. stomach, m.t. Malpighian tubules, mb. intestine, 7.g/d. rectal glands.
Some of the crop-contents of G. tetraonis, pressed out. It consists of
feathers in various stages of disintegration.
Egg of Menopon pallescens Nitzsch. Under pressure the operculum has
come off and the ovum is squeezing its way out.
Four eggs of Goniodes tetraonis, attached to the base of an after-plume,
between it and the shaft of the plume of a feather from the flank. In one
of them the operculum has fallen off.
Pratt XLI.
. Optical section of abdomen of G. tetraonis, showing a. anus, c. crop, fb.
fat-bodies, int. mtestine, 7.glds. rectal glands, ov. ovary.
. Nirmus cameratus Nitzsch. Dorsal view of female.
. Ventral view of the same.
Prate XLII.
. Ventral view of head of Nirmus cameratus, showing mouth-parts, antenne,
eyes, position of anterior legs.
. Eggs of Nirmus cameratus on the feathers of a young grouse approxi-
mately three weeks old. A. Very slightly magnified; three eggs on one
of the wing-coverts. . Magnified about eight times on a downy plume.
C. Very highly magnified to show the reticulations.
Prare XLII.
. Ornithomyia lagopodis Sharp. A. dorsal view, B. ventral view, C. dorso-
lateral view of toot.
Pratt XLIV.
A ventral view of the anterior edge of the head of O. lagopodis and the
mouth-parts. Ant.antenne, e. eyes, map. maxillary palp, prob. proboscis.
A still more enlarged view of the orifice of the proboscis is shown to the
right.
23. A figure of part of the femur, f.; the tibia, ¢7., the tarsus, ¢a., and the foot
of O. lagopodis.
. The same, more highly magnified, from the inner or under surface. emp.
empodium, ped. pulvillus.
. Five sketches of the pupa of O.lagopodis. A. Life-size figure of the mature
pupa; B. the same of an immature pupa dissected out of a fly; C. mature
pupa-case magnified and seen from the micropyle end; D. a similar view
of the immature pupa; E. mature pupa-case seen from above; F. the
same seen from the side. m. micropyle, m.a. micropylar area.
Prate XLV.
. The larva of Scatophaga stercoraria L. A. Life-size: B. magnified. a.s.
pag g
anterior spiracle, p.s. posterior spiracle, h.s. hypostomal sclerite, m.s.
mandibular sclerite, ph.s. pharyngeal sclerite, v.¢. visceral trachea,
¢.c. transverse commissure.
. Enlarged view of oral and pharyngeal sclerites, lettering as in fig.26. d.g.,
dental sclerite.
PrarE XLVI.
. Exlargec view of head of the maggot, showing a.s. anterior spiracles,
spines, more pronounced at the anterior end of each segment, m. mouth,
f.b. fat-body ; other lettering as in fig. 26.
29. Lateral view of mouth of the maggot, showing sensory papillae. Magnified.
30. Dorsal view of the head of the maggot, showing the anterior spiracles
protruded. Magnified.
31. Dorsal view of posterior end of maggot, showing p.s. posterior spiracles, the
split tracheal main trunks and papillae. Magnified.
. Lateral view of the same, showing a. the anus. The specimen was slightly
compressed and the posterior spiracles were a little displaced and both
brought into view.
Pirate XLVII.
Side view of male specimen of Ceratophyllus gallinule Dale.
Side view of female specimen of the same drawn to scale.
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TRIGHOSOMA LONGICOLLE.
CONTENTS OF ALIMENTARY CANAL.
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1909. } ON THREAD-WORMS OF THE RED GROUSE. 399
2. The Thread-Worms (Nematoda) of the Red Grouse
(Lagopus scoticus). By A. E. Surpiey, M.A., Hon.
D.Sc., F.R.S., F.Z.S., Fellow and Tutor of Christ's
College, Cambridge, and Reader in Zoology in the
University.
[Received December 12, 1908.)
(Plates XLVIIL-LV.*)
NEMATODA. Thread-worms or Round-worms.
(i.) Family Strongylide.
(i.) TRICHOSTRONGYLUS PERGRACILIS (Cobbold).
Synonym: Strongylus gracilis Cobbold.
History.
This round-worm was first described under the name of Stron-
gylus pergracilis Cobbold, by Cobbold + whose words we quote :—
“ Characters.— Body filiform, finely striated, gradually dimi-
nishing in front, uniform in thickness below; head bluntly
pointed, with a simple oral aperture; tail of the male furnished
with a bilobed bursa, each half supporting four pointed rays;
spicules two, thick, and slightly divergent; tail of the female
slightly swollen above the subterminal anal orifice, rather sharply
pointed at the tip; vaginal opening situated at the upper part of
the inferior sixth of the body.
“Length of male 3" to 2”; body =4," in diameter, tapering
anteriorly to Z000, at the head; greatest breadth immediately
above the bursa ashy:
“ Length of female mostly 2 2 sometimes very nearly 3”
breadth above = tail 4,” to ;4,’, narrowing at the extreme
point to —),,’ ieee diameter of the eggs 1,", their
breadth being =1,"'.”
Eight years later Cobbold described, under the name of
S. douglassii, a nematode which occurred in great numbers in the
proventriculus of certain South African ostriches. Their presence
was associated by the ostrich-farmers with a certain amount of
disease and with some deaths.
Two other allied forms, Trichostrongylus (Strongylus) nodularis
and Trichostrongylus (Strong ylus) tenuis, are also held to cause
disease in birds, whilst recently a fifth form, S. quadriradiatus
(possibly also a Trichostrongylus), has been added by Stevenson $
to the list of the Strongylids harmful to birds.
id
* For explanation of the Plates see P. 349.
Mee ‘The Grouse Disease,’ ‘ The Field ’ Office, London, 1873.
{ J. Linn. Soc. London, Zool. xvi. 1883, p. 184.
§ U.S. Dept. Agric., Bureau Anim. Industry, Cirenlar 47, 1904.
336 DR. A. E, SHIPLEY ON [ Mar. 16,
T. nodularis* Rud. is found in the mucous and muscular coats
of the stomach and duodenum of many species of Anatide. It.
has frequently been observed in the domestic goose and is especially
fatal to the young of this bird.
T. tenuis Eberth ft occurs in the cecum of the grey goose,
Anser cinereus Meyer, and of the domestic goose. Neumann
states that it is “‘ rare.”
Finally, we have the species S. quadriradiatus recently described
by E. C. Stevenson. It occurred in considerable numbers in the
intestines of a flock of fancy pigeons which had been almost
destroyed by a malady of unknown origin early in 1904. In his
article upon this epizootic, Stevenson points out that the presence
of a few nematodes in the cecum of the pigeon causes little harm.
If, however, the thread-worms exist in large numbers, disease
becomes manifest. This Stevenson attributes to two causes: the
first is the loss of blood ; but there is, I think, little or no evidence
that these nematodes live on blood. The second cause is the
piercing of the walls of the intestine t, which permit the bacteria
of the contents of the alimentary canal to make their way into the
peritoneal cavity, where they set up peritonitis. Evidence is
gradually accumulating as to the occurrence of this, and some of
the French authorities even think that such a perforation, made
as a rule by 7richocephalus dispar, is one of the more common, if
not the most common, causes of appendicitisin man. The presence
of these worms further sets up an inflamed, catarrhal condition
of the walls of the intestine, which leads toa debilitating diarrhea
and to general disorders of the digestive system. As in other
cases, the Nematodes doubtless give off toxins, the effect of which is:
largely confined to the nervous system and to the cells in the
blood of the host.
SrRUCTURE.
The genus Trichostrongylus has recently been established by
Looss § to include certain forms which he has separated out from
the large genus Strongylus. The Strongylus pergracilis of Cobbold
corresponds so closely in structure with the species described by
Looss that I think there can be no doubt that it also should be
included in the new genus. The suggestion made above that
Strongylus tenuis Eberth of the goose should also be reckoned
as a species of Z'richostrongylus was made to me by Mr. R. T.
Leiper.
Specimens of 7’. pergracilis are found in the cxca of grouse.
They are apt to cover themselves with mucus and dirt, and are
* Wedl, SB. Akad. Wien, xix. p. 46, and Schneider, Monogr. d. Nematoden,
p. 148. Nan
+ Wiirzburger nat. Zeitung, ii. p. 47. Von Linstow, Troeschel’s Archiv, 1. 1876,
5 Gb
{ An actual perforation of the membrane is not in all cases necessary. There are
examples of bacteria traversing the wall or parts of the wall of the alimentary canal
which have been locally or temporarily weakened in some way.
§ Centrbl. Bakter. Parasitenk. xxxix. 1905, p. 409.
1909. | THREAD-WORMS OF THE RED GROUSE. 337
consequently hard to see and often overlooked. We have found
them, with hardly an exception, in every one of the hundreds of
grouse examined. They are rendered opaque and white, and
hence much more apparent, by shaking up the contents of the cecum
in 75 per cent. alcohol, to which a few drops of corrosive sub-
limate have been added. Their presence is also readily detected
by compressing a drop or two of the cecal contents between two
microscope-slides and holding them up to the ight. The worms,
if there be any, then appear as thin, white, transparent lines.
We owe this method to Dr. Wilson.
T. pergracilis is an extremely fine worm, measuring in the male
on the average 6 to 8 mm. and in the female 8 to 10 mm.
They are very narrow and hair-like and, as a rule, whitish in
colour, but sometimes have the tinge of blood when seen in a
very thin layer on a slide through the microscope. ‘They are very
transparent, readily revealing their internal structure, and they
are so soft that the pressure of a cover- shp almost always ruptures
them. The cuticle is very clearly and definitely ringed ( Pl XLT,
figs. 3 & 5), and the rings are so constituted that whilst the worm
can easily work its way forward through a tissue, it would have
difficulty in wriggling backward. The rings give the edge of the
body a strongly serrated appearance like a saw. ‘This is most
marked a little way behind the head and extends over about cne-
third the body length. There is no trace of longitudinal marking
on the cuticle.
The genital bursa in the male is well formed, and opens to the
exterior by an oval opening with its long axis longitudinal. The
bursa 1s supported by a number of ridges as an umbrella is by its
ribs, and, using Looss’s nomenclature, these are arranged in three
groups. The members of each group arise from a common root
and are recognisable, even when, as in the case of 7’. pergracilis,
some of them run close to and parallel with members of another
group. The three groupsare: (1) Dorsal, (2) Lateral, (3) Ventral.
The Dorsal group consists of a median, posterior dorsal rib,
which is forked slightly at its free end ; oer the same root with
it arise a pair of externo-dorsal ribs, one of which passes into the
side membrane of the bursa (Pl. L. figs. 11 & 12). In front of the
externo-dorsal, three lateral ribs arise from a common root ; on each
side of these the most posterior or postero-lateral vib is smaller than
the others and slightly separated from them; the median-lateral
rib and the antero-lateral rib are of similar size and lie parallel.
There are on each side in front of the lateral ribs a pair of ventral
ribs; of these the hindermost or latero-ventral rib forsakes its
group and comes to lie close and parallel to the antero-lateral rib,
which it further resembles in size and shape. The ventro-ventral,
on the other hand, is simmer and stretches forward to support.
the anterior edge of the bursa. The whole arrangement of the
lateral and ventral ribs recalls a slightly flexed hand, with the
thumb thin and extended and the little finger slightly divaricated.
The thumb represents the ventro-ventral rib, the first finger the
338 DR. A. E, SHIPLEY ON [ Mar. 16,
latero-ventral rib, the second finger the antero-lateral rib, the
third finger the medio-lateral rib, and the little finger the postero-
lateral rib (PJ. XLIX. fig. 10).
The spicules in the male are very conspicuous and very difficult
to describe. Figures 11, 12, 13 on Pl. L. give a better idea of their
appearance than any verbal description can. They are short,
strongly chitinised, with thickened edges and a kind of haft or base
at the anterior end; and each spicule is hollowed something like
a crumpled, withered, lanceolate leaf. Each spicule is provided
with retractor and protractor muscles, and, as fig. 12 shows, when
protracted they are divaricated. When in this extruded condition
they form a cross, the left spicule projecting to the right and
vice versd. Besides the spicules and between them, rather to the
posterior end, lies the accessory or median piece, which Looss calls
the “gubernaculum.” It is best seen in profile, and has then
somewhat the outline of a Turkish slipper. It also has muscles
inserted into its ends.
Near the base of each spicule is an oval clear vesicle, but
apparently the end of the spicule was outside and not inside the
lumen of the vesicle.
The head presents very little signs of differentiation (Pl. X LVI.
fig. 3). In some specimens with a one-twelfth objective three very
minute lobes can be seen, but they are not visible in all cases and
their appearance may be due to some expansion of the mouth. The
mouth is terminal and leads into a slightly bulb-like cavity which
soon narrows into the thin capillary lumen of the alimentary canal.
The cesophagus is more granular than the intestine, and separated
from it by a very shallow groove ; its walls consist of flatly rounded
cells with conspicuous nuclei (fig. 5). I could not detect any parts
in the intestine ; it appears to be an undifferentiated tube running
from mouth to anus, the lumen lined with chitin and the walls
formed of granular cells with visible nuclei. No food was seen
in the alimentary canal. Posteriorly the intestine widens into a
spacious rectum, which just in front of the anus narrows again
into a short, thin, terminal portion. The anus in the male opens
into the genital bursa; in the female it isa little distance in
front of the end of the pointed tail, but relatively net so far forward
as it is in the larva. Two cervical glands run about a fifth of the
length of the body backwards, and end with rounded ends about
the same level.
In the male the testis begins about the level where these glands
end. It consists of a single tube, the cells lining which give
rise to the spermatozoa. Anteriorly the cells when squeezed out
seem amceboid, with rounded and very refringent nuclei. The
hinder end of the testis is, however, crowded with spermatozoa
shaped like little squat bottles, and in some specimens the genital
bursa sheltered two clumps of these, looking as though they had
escaped from two vesicule seminales.
I saw nothing of excretory canals or their opening, and unless
an ill-defined ring which surrounded the alimentary canal, about
1909. ] THREAD-WORMS OF THE RED GROUSE. 339
one-twenty-fifth of the body-length from the anterior end, be the
nerve-ring, I saw nothing of the nervous system.
The ovaries are double. Hach tube arises about the level or a
little behind the level of the hinder end of the cervical glands.
One of them runs, with but slight undulations, straight to the
“ovejector”” which opens by the vagina, situated about one-sixth
of the body-length from the hinder end, the other passes the vagina
and reaches back almost to the anus; it then doubles forward
again and opens into the posterior ‘“ ovejector.”
The anterior end of each ovary contains undifferentiated
protoplasm, but soon eggs begin toappear. At first these are very
flattened, like a pile of coins, much broader than they are long;
then they become thicker, and, finally, three or four times longer
than they are broad. The rounded nucleus is in every stage
very conspicuous. It is impossible to say precisely where the
ovary ceases and the oviduct begins. We find the long cylindrical
cells rounding themselves off and an egg-shell beginning to appear.
By this time fertilization must have taken place, but I have not
seen any spermatozoa in the oviduct. The oval cells usually he
at first with their long axis at right angles to the longitudinal
axis of the oviduct; then, when a little older, they lie obliquely, and,
finally, they come to lie with their long axes parallel to that of the
duct, in which position they are most readily swallowed by the
‘“‘ovejector.” The anterior and the posterior oviducts usually
contain one, two, or three unsegmented ova; then come some six
to eight segmented eges representing as a rule the stages with two,
four, eight, sixteen, thirty-two, sixty-four, and sometimes a hundred
and twenty-eight blastomeres. One or two of these stages may be
represented by two ova, but in any case the segmentation must
be very rapid.
The lower end of the oviduct is lined by what in optical section
appear to be high columnar cells with very granular disintegrating
borders. These seem to be secreting something. The walls of
the oviduct pass suddenly into the ‘‘ ovejector,” which consists of
three parts: (a) The most internal is somewhat funnel- or trumpet-
shaped, its wider mouth is continuous with the walls of the oviduct
and is crenellated ; the funnel is richly supplied with both longi-
tudinal and oinetler fibres; not infrequently it contains an egg.
(6) The second chamber of the “ovejector” is spherical, very trans-
parent, and is marked by the presence of a large number of radiating
muscle-fibres running from the periphery to the limits of the lumen.
The contraction of these fibres would enlarge the lumen and suck
the egg on. (c) The third chamber of the ‘“ovejector” is thin-
railed, with a chitinous lining. It frequently shelters an egg.
At its outer end it narrows, and uniting with the similar narrow
end of its fellow it forms an extremely short vagina which opens
to the exterior by a longitudinal slit, the edges of which are
also crenellated.
The ova are laid in the fluid contents of the host’s ceca, in
which they are frequently found floating. We have found
340 DR. A. E. SHIPLEY ON _ [Mar. 16,
developing ova in the ceca of a young grouse chick of 7 to 10
days of age from Auchenterlie, Dumbar tonshire. Apparently the
ceca are the chief centres of absorption of the digested food ;
they contain none of the cellulose skeletons of vegetable cells so
common in the intestine, and none of the masses of cast epithelium
which make up so large a proportion of the flocculent masses in
the duodenum. The eges may develop further inside the cecum,
though as yet we have not found an egg containing an embryo in
its contents.
A small pellet of the cecal contents, such as can be carried
away on the point of a needie, spr ead out under a cover-slip,,
will, in a well- infected bird, show some 12-20 worms and 100-200
eggs in the field of a § 2 Ross’ objective with a No. 2 eyepiece.
ALLIED SPECIES.
In his memoir on the genus Trichostrongylus, Looss enumerates.
the following four species :—
(1) 7. rerorr«roruis (Zeder), 1800. From the duodenum
and exceptionally from the stomach of Lepus timidus and Lepus
cuniculus (when undomesticated). Railliet says it coexists with
Strongylus strigosus, and helps to give rise toa pernicious anzemia..
It develops directly without intermediate host.
(2) 2. msrapreis (Railliet), 1893. Syn. 7. subtilis Looss,
1905. From the duodenum and exceptionally from the stomach of
Ovis aries, Ovis laticauda, Antilope dorcas, Camelus dromedarius:
(Egypt), Papio (Cynocephalus) hamadryas (North Africa), and
occasionally in Man (Egypt and Japan). RRailliet states that this
species, together with Strongylus contortus, lives in the duodenum
of sheep, which succumb to pernicious anzemia.
(3) 7. prozwoLurus (Railliet), 1896. From the duodenum of
Ovis aries, Ovis laticeuda, Antilope dorcas, and occasionally of
Man (Egypt), and Camelus dromedarius (¢ Paris and Egy pt).
(4) Z. rizrrnus Looss, 1905. From the duodenum of Ovis aries,
Ovis laticauda, occasionally from Camelus dromedarius and Man
(Egypt). Looss regards this as a rare species.
To these must be probably added :—
(5) 7. rewurs (Kberth), 1861. Syn. S. tenwis (Mehlis) Eberth,,
1861. From the cecum of the goose, Anser cinerea, and
(6) Z. woputaris (Rud.), 1809. Syn. S. nodularis Rud.,.
1809; Ascaris mucronata Frohlich, 1791; S. anseris Zeder, 1800 ;
S. nodulosus Rud., 1803; S. crispinws Molin, 1850. From the-
mucous and muscular coats of the stomach and duodenum of
various species of the family Anatide. They are said to be very
fatal to young geese.
(7) 7. Percracizis (Cobbold), 1873. Syn. S. pergracilis-
Cobbold, 1873. From the cecum of Lagopus scoticus.
(8) 7’. QUADRIRADIATUS (Stevenson), 1904. Syn. S. quadri--
radiatus Stevenson, From the intestines of pigeons.
1909. | _ 'THREAD-WORMS OF THE RED GROUSE. 341
(9) 7. exrenwaTvs (Railliet), 1898. Syn. S. gracilis McFad-
yean, 1897, not Leuckart, 1842. This form occurs in the fourth
stomach of cattle in England, and in cattle, sheep, and goats in
the United States *.
(10) 7. capricora Ransom, 1907. From goats and sheep in
the United States.
It is noticeable that the parasite occurs only in vegetable
feeders, and that, whereas it lives always in the stomach or the
duodenum of Mammals, it chiefly inhabits the ceca of Birds.
PARASItEs OF 7'RICHOSTRONGYLUS PERGRACILIS (Cobbold).
Ina female specimen two ameeboid organisms were making
their way along the body-cavity in the region of the “ ovejector.”
Each was throwing out rounded pseudopodia, and the distinction
between the granular endosare and the glassy ectosare was very
sharp. Another specimen had some refringent bodies, in shape
like short rows of yeast-cells or fungi-spores, lying in the body-
eavity (Pl. LITT. fig. 32).
Lire-Hisrory oF U'RICHOSTRONGYLUS PERGRACILIS (Cobbold).
The eggs are found in all the earlier phases of segmentation in
the fluid contained in the intestinal czeca of the grouse, and usually
attain there a stage of 32 or 64 blastomeres. Removed from the
body and kept in a watch-glass in water at the temperature of a
warm room they continue to segment until in the course of
twenty-four hours the larve, typically coiled in the egg-shell
(Pl. LI. fig. 23), are found. A shght pressure on the egg-shell
causes a rupture, the shell instantly contracts (compare fig. 24
with fig. 23 on PJ. LI.), and the larva begins to come out.
Exactly similar stages of development are found in the feces
of the grouse, both in the rectum and after leaving the body, and
the ova containing the coiled embryos of the Trichostrongylus are
common in grouse-droppings. A microscopic examination of
these droppings reveals a wonderful collection of objects. In one
specimen alone we found (1) moss-spores and undigested moss-
capsules, (2) bits of Spirogyra, (3) diatoms, (4) the hyphee of
some fungus, (5) undigested epidermal cells showing stomata
from a leaf, (6) spiral woody fibres, (7) bacteria, (8) portions of
feathers, (9) the leg of a spider, (10) specimens of dAmeba or of
ameeboid organisms, (11) cestode eggs, (12) eggs and the head-
end of Trichosoma longicolle Rud., (13) eggs with larva of Z’richo-
strongylus pergracilis (Cobbold), (14) erystals which are the same
as those which abound in the ceca. Two kinds of these are
abundant, one resembling the Charcot-Leiden crystals, the presence
of which is usually associated with parasites in the intestine,
the other resembling the crystals of magnesium ammonium
phosphate, Mg(NH,)(PO,), 6H,O. Doubtless further search would
* B. H. Ransom, U.S. Dept. Agric., Bureau Anim. Industry, Circular 116, 1907.
342 DR. A. E. SHIPLEY ON | Mar. 16,
result in the discovery of many other objects. The important.
fact from our point of view is that the expelled feces contain
ova within which are fully formed larve ripe for hatching.
But we have also found hatched larvee in the feces which were
still in the rectum, so we may conclude that the droppings may
contain (1) larve coiled in the egg-shell, and (2) larve which
have escaped from the egg-shell and are living freely.
No one has ever seen, and probably no one will ever actually
see, one of these eggs or larve of the thread-worm enter the body
of the grouse, but that they do so directly is, I think, unquestion-
able. Railliet* and Ransom tT have both succeeded in infecting
rabbits by directly feeding them on the embryos of 7. retorte-
formis ; and presumably the development of 7. pergracilis is also,
like that of its congener, accomplished without an intermediate:
host. In Ransom’s case the embryos apparently hatched out and
developed to the ensheathed stage within two weeks after the
passage of the eggs from the intestine of the host, and the
embryos when swallowed by a second rabbit became mature and
capable of laying eggs in about a month after they had been
‘swallowed. The thread-worms exist in hundreds in every bird ;
the female thread-worms, which greatly exceed the male thread-
worms in number, lay thousands of eggs. The eggs and larvee
exist in countless numbers in all grouse-droppings. As these are
disintegrated by rain and frost, these minute organisms are either
washed by the water on to all the food or pebbles a grouse may
pick up, or in the absence of rain are dried and blown by the
wind all amongst the heather and other food which a grouse may
eat. The ova and larve of thread-worms are notoriously resistant
to the effects of desiccation, so that even in the event of prolonged
droughts—which, judging from my experience of Scotland, must
be a remote contingency—the eggs and larve are in no way
prevented from reaching their host <.
During a visit to Beaufort Castle in August, 1908, I introduced
what has since, in the hands of Dr. E. A. Wilson, proved a
valuable instrument in research. We had hitherto been unable
to find the ova or the young nematodes upon the heather; they
are so minute and require so high a power of the microscope to
see them, that all attempts to look at them on the stalks or in
the buds had failed. The new method showed that nematodes,
some of them free-living, and their larve and the larve of
parasitic forms exist in countless thousands im every square foot
of heather.
The method consists in soaking a handful of heather in a glass
bottle, using just as little water as will well cover the twigs, for
several hours—overnight is not too long; then shaking the bottle
well for some minutes, or shaking it more gently in a rocker
* Bull. Soc. Zool. France, xiv. p. 375.
+ U.S. Dept. Agric., Bureau Anim. Industry, Circular 116, 1907.
~ Since the above was written, Mr. R. T. Leiper has obtained all the stages of the
life-history of 7. pergracilis and has succeeded in infecting hand-reared grouse
which were previously free from the worms.
1909. ] THREAL- YORMS OF THE RED GROUSE. 343:
for a longer period, and allowing the sediment to settle. The
sediment is then, with as little water as possible, taken up
by a pipette and placed in the test-tube of a centrifugal machine.
After a few minutes’ rapid rotation the heavy matter which
accumulates at the closed end of the test-tube is removed and
examined microscopically. The very first time we tried this
method we succeeded in showing that heather is, so to speak,
“crawling” with thread-worms, amongst which we thought we
recognised the larvee of 7’. pergracilis. We also found aphides, a
dipterous larva, thousands of pollen-grains, tardigrades, thysanura,
rotifers, nematodes, vegetable débris, desmids, beetle-larvee,
young hemiptera. One of the nematodes had striated cuticle
and was about the proportions of 7. pergracilis; some of the
others were relatively much longer and seemed to be free-living.
Amongst the nematodes was a male with the characteristic
spicules of the 7. pergracilis, of the same sort of outline
and in the same relative position. The tail was curved and
showed as yét no genital bursa. The mouth led into a short
cesophagus (| ) with a small swelling. The body contained a.
eranular substance.
These eggs and larve must exist in quite countless millions,
every dropping must contain many thousands; roughly speaking,
a grouse—at any rate during the night—relieves itself about once
an hour, and the droppings occur every few yards on the moors.
Hence the occurrence of Trichostrongylus pergracilis even in
grouse chicks of a tender age is readily explained.
One must not forget that a grouse-dropping consists of two
parts: (1) the excreta from the intestine, which contains the tape-
worm and 7'richosoma eggs; and (2) the more fluid contents of the
ceca, Which alone contains the Vrichostrongylus eggs and larve.
This excretion usually forms a cap upon the firmer dejecta from
the intestine, but sometimes occurs alone ; it either case it probably
is washed away by the first rain or quickly dries up, whilst the
more solid excreta will retain their shape for a couple of years
before disintegrating, as is shown by finding them unchanged in
shape but with charred ends on moors that were burnt some two
years ago.
We have seen that a young grouse-chick, not more than ten
days old, contains in its ceca segmenting ova, young larve, and
adult males and females, and we are faced with the old problem
as to whether the egg preceded the fowl or the fowl the egg.
Are the segmenting ova we find in these young ceca of the same
generation as the larvee and adults, or are they the offspring of
the adults¢ On the whole, I incline to the second alternative.
I think most probably that the larvee, either coiled up in the ege-
shell or free, obtain access to the alimentary canal with the food
or water, make their way to the ceca, and there, remote from the
world, in the warmth and obscurity of their narrow home, rapidly
mature, couple, and quickly begin to lay eggs, and that these
344 DR. A. E, SHIPLEY ON [ Mar. 16,
eges are those which we find in the ceca of the young grouse-
chick.
This, then, appears to be the ordinary life-history of Z'richo-
strongylus pergracilis, but it is sometimes varied, and varied in a
way which may have a bearing upon grouse-disease. We have
seen that as a rule eggs with coiled-up larve leave the body of
the bird, but we have also seen that free larvee have sometimes
been found in the feces whilst still in the rectum. We have
further discovered small nematode larve in the contents of the
small intestine. These larve are more pointed than the larvee
we have watched hatch out from the Trichostrongylus eges;
still they may be the larve of this species*. Dr. Wilson has
further found on one or two occasions both in the lungs and in
the liver certain larval forms which resemble the larve we
have artificially hatched out of the developing eggs of J. per-
gracilis, and the larvee which we have found in the dejecta of the
grouse.
These larve are extraordinarily minute, and unless seen alive
are very difficult to recognise even under a twelfth oil-immersion.
Hence they have hitherto been overlooked by researchers who
have necessarily had to deal with grouse that had been dead for
many hours, often for days. If these larvee be in truth the young
forms of 7’. pergracilis, then under certain—at present undeter-
mined—conditions, they either proprio motu or carried by the blood-
stream reach the two important organs, the liver and the lungs.
The cavity of the alimentary canal is infested with bacteria, all
very well in their place, but liable to become pathogenic if carried
to other tissues and organs. It is improbable that these larvee
with their smooth, dry, ‘‘ polished ivory” surface carry such
bacteria adherent to their outside, but it is quite possible that
feeding as they do on the fluid contents of the digestive tract
they ingest such bacteria, and during their ‘‘ Wanderjahre ” deposit
them undigested in just those organs where their pathogenic
properties are most easily stimulated.
Apart from the harm thus caused, the presence of the Tvicho-
strongylus produces grave local troubles, such as inflammation,
peritonitis, ulceration, &e., which will be dealt with by the patho-
logists now working for the Inquiry.
Gi.) SYNGAMUS TRACHEALIS von Sieb.
The Red or Forked-Worm.
We have found this common pest of the fowl-yard and
heasant-coop but twice in the grouse. Probably their free and
unconfined life, together with the comparative paucity of earth-
* It is also possible, and perhaps more probable, that they are the young of
Trichosoma longicolle Rud.
1909. | THREAD-WORMS OF THE RED GROUSE. 345.
worms in the moors, protects grouse from the ‘ gapes,” as the
disease caused by the forked-worm is called. Earthworms abound
in Scotland in the cultivated lands, pastures, and woodlands, and
occur even on the tops of mountains. Mr.:-Wm. Evans tells me
he has a list of seventeen species of the Lumbricide taken north
of the Tweed, but they are practically absent from the peat-moors,
where heather, grouse, and humic acid are most abundant. Some
moors, however, such as those on the Pentlands, include patches
of land in which worms flourish. Still there seems ne reason at
present to incriminate the forked-worm of causing any trouble
to grouse. One of our cases was a young bird from Argyllshire.
Gi.) Family Trichotrachelide.
Gu.) Z'RICHOSOMA LONGICOLLE Rud.
Synonyms: Calodiwm caudinflatwm Molin.
Trichosoma gallinum Kow.
Trichosoma caudinflatum Kow.
History.
This genus, first named and described by Rudolphi*, is also
described by Dujardin? from various species of gallinaceous
birds. He gives a list of the earlier and ill-defined synonyms.
The species is also mentioned by Diesing £ under the same name.
It is recorded by Molin § under the name Calodiwm caudinflatum,
the name referring to the swollen tail of the female, from the
small intestine of the partridge and the quail. Eberth ||, who
gives the best account of the anatomy of the genus TJrichosoma,
gives a short description and a figure of a nematode under the
same name, Z'richosoma longicolle; he draws attention to Du-
jardin’s description of a funnel-lke appendix to the vagina and
surmises that this is a prolapsed piece of the vagina; this is
undoubtedly the case in some of our specimens (Pl. LIT. fig. 29, ii.).
Kowalevsky 4 mentions under the name Zrichosoma gallinwn
what Dr. von Linstow tells me is this species, and in a later
paper ** apparently describes and figures this under the name
Trichosoma caudinflatum. Untortunately I have been unable to
read this, as the paper is written in Polish. Railliet +? considers
that the Zrichosoma longicolle described by Dujardin’ and Eberth
does not agree with Rudolphi’s description of his ZVrichosoma
longicolle. He points out that Dujardin and Eberth’s species
* C. A. Rudolphi, ‘ Entozoorum Synopsis,’ Berlin, 1819.
+ M. F. Dujardin, * Hist. Nat. des Helminthes,’ p. 19.
+ C. M. Diesing, ‘Systema Helminthum,’ 1850-41, ii. p. 260.
§ R. Molin, 8.B. Ak. Wien, xxxiii. 1859, p. 302, and Denk. Ak. Wien, xix. 1861,
. 330, also Mem. Ist. Veneto, 1x. 1860, p. 617.
|| C. J. Eberth, ‘ Untersuchungen tiber Nematoden,’ Leipzig, 1863.
@ M. Kowalevsky, Bull. Ac. Cracovie, 1894, p. 280.
** M. Kowalevsky, Rospr. Ak. Krakov. xxxviil. 1901, p. 274.
++ A. Railliet, ‘Traité de Zoologie Médicale et Agricole,’ Paris, 1895.
Proc. Zoou. Soc.—1909, No. XXIII. 23
346 DR. A. E. SHIPLEY ON [ Mar. 16,
occurs in the caecum of the fowl and of the guinea-fowl, and
he renames this species Zrichosoma retuswm Raill., 1893. The
length of this worm is 13 mm. in the male, 19 mm. in the female.
Rudolphi’s worms—which may belong to more than one species—
vary from 39 to 80 mm., and have been described from Lyrurus
(Tetrao) tetrix, the Black Grouse, Black-cock, or Grey-hen ;
Tetrao wrogallus, the Caperecaillie; Gallus gallinaceus, the Common
Fowl; fhasianus colchicus, the Common Pheasant; Ohryso-
lophus (Phasianus) pictus, the Golden Pheasant; Perdix cinerea,
the Common Partridge; and Coturnix communis, the Common
Quail.
We first found specimens of 7’. longicolle in a Perthshire grouse
which was brought us in the morning we were leaving Blair
Atholl for the south in the autumn of 1906. Having once
seen it, however, it was soon observed again, though it occurs
sparingly. It always lives in the duodenum, sometimes associated
with the tape-worm Hymenolepis nicrops, and sometimes alone.
‘The worms resemble short pieces of very fine white silk.
This species has two longitudinal rows of dark spots irregularly
scattered in two lateral bands. Roughly speaking, there are five
or six of the spots in a transverse row, but they are not regularly
arranged. The two bands arise anteriorly in the region of the
cesophagus, and as they pass backward they become somewhat
narrower, much more pronounced in appearance, and darker.
They end on the extreme end of the body (Pl. LIV. fig. 36). Hach
spot corresponds with a unicellular gland, and the bands of these
elands replace the ordinary nematode excretory system in the
Trichotrachelide, the family to which 7richosoma belongs. They
have been best described by Jiigerskidld*. Hach cell opens by a
minute straight duct which traverses the cuticle and forms what
used to be called the rod-shaped body (Pl. LIV. fig. 37). The
trichosoma longicolle of Eberth + has a third or ventral band, and
he mentions that Dujardin saw but one band in his specimens.
The length of the specimens varied from 20 mm. in the male
to 40 mm. in the female. The greatest breadth of the body
was 4:5 4, but in the neck-region it did not exceed 3p, and
tapered away to the anterior end, where the breadth was but 0°5 p.
The very regular large cells in the region of the neck, which
are pierced by the cesophagus, are just under 3 in width and
are 12y in length. In the youmg specimens these cylindrical
cells with flat ends, lying like a lot of pillars end to end, are not
cut up into a series of segments, which gives a scolloped outline
to the cells of the adult when seen in profile. But latera number
of circular constrictions arise, and these divide each cell into a
series of ten or twelve areas upon each side, and the whole cell
has the appearance of being built up of two rows of rounded
bricks lying side by side in a double pile. The nucleus remains
large, oval, and conspicuous. At the end of each cell there is
* Svenska Ak. Handl. xxxvy. 11. (1901).
+ ‘Untersuchungen tiber Nematoden,’ Leipzig, 1863.
1909. ] THREAD-WORMS OF THE RED GROUSE. 347
usually a dark granulation which serves very clearly to define
their limits (Pl. LIV. fig. 35).
The lumen of the cesophagus which pierces these peculiar cells
is very minute, and is lined with a definite chitinous tube, The
“cellular body,” as the aggregate of the cesophageal cells is some-
times called, ends abruptly, about one-fifth the body-length from
the anterior end. Here the cesophagus passes quite abruptly into
the capacious intestine with its many-celled wails. Just at this
point, and tucked away in the angle formed by the minute
esophagus widening into the broad intestine, are a couple of
glands, probably the homologues of the cervical glands of other
nematodes (Pl. LIV. fig. 38). The intestine continues to the
hinder end of the body with no change; it is somewhat difficult
to distinguish, as it is just about the same brown colour as the
lateral lines. The posterior end of the female is truncated and
the anus is at the ventral side of the abruptly terminated body.
The ovary is a single tube which anteriorly contains undif-
ferentiated eggs. These gradually attain a definite and somewhat
irregular outhne. Posteriorly the ovary opens into a spacious
uterus, in which the ova are oval, with a distinct vitelline mem-
brane. The uterus is broad, and serves, with its contents, to
conceal the other organs of the body. Posteriorly, where the
body is wide, the ova are irregularly crowded together; there
may be as many as five or six ova in a transverse row. Further
towards the head the ova acquire their characteristic egg-shell
with two bright spotsateach end. They closely resemble the eggs
of Trichocephalus trichiurus (dispar) (.) (Pl. LIV. figs. 40 & 41).
About halfway along the body the diameter lessens as we pass
forward, and after a certain space the uterus is narrowed and only
permits a couple of eggs to be abreast, and finally the eggs are
reduced to a single row. ‘The uterus opens by a vagina which is
situated a little way behind the end of the “ cellular body,” ¢.e. just
behind the anterior end of the intestine. The uterus or vagina
is usually prolapsed and forms a bell-like structure, one edge
of which usually has a clear oval vesicle in its walls (BE Tein
fig. 38). Through this bell-shaped structure the eggs pass out.
“The male is markedly smaller than the female, Its average
length is about 25 mm. and its width throughout does not surpass
the anterior end of the female’s body.
The testis is a single tube which opens posteriorly. At the
tail end the male has a pair of cuticular folds or flanges, possibly
representing a genital bursa. There is a single spicule, very
long, and in many cases only perhaps protruded for a fourth or
fifth of its whole length. It is described as having a sheath, but
in the specimens we have seen this was not appar ent—probably it
was retracted (Pl. LIV. fig. 39).
The males are very much rarer than the females—in fact, we
examined a considerable number of specimens without finding a
single male, probably they occur in about the proportion of one
to seven or ten females. We have occasionally found a very
23*
348 DR. A. E. SHIPLEY ON [| Mar. 16,
long, thin larva in the duodenum, which we take to be the larva
of tie Trichosome.
The eggs appear to undergo no segmentation in the body of
the worm, and, in fact, we have not yet seen an egg of Trichosoma
longicolle segmenting. In one grouse from Ross-shire small
embryos of some nematode were found in the small intestine.
It is possible that these are the young of 7. longicolle, but
they show no trace of division into neck and body. It is also
possible that they are the larve of Zvrichostrongylus pergracilis,
but they differ in size and shape from those young of this species
which we have hatched out and found free. The grouse in which
they were found had been feeding on corn, and J am rather
inclined to believe that these larve are the young forms of
Tylenchus tritici which causes the well-known corn-cockle.
Trichosoma longicolle occurs only in the duodenum, often
associated with the species of Hymenolepis which inhabits this
part of the alimentary canal. They are surrounded by epithelial
cells, singly and in clumps, and of many sizes and shapes, which
have been shed in immense numbers from the wall of the
duodenum (Pl. LV. figs. 42 & 43). These may be detached by post-
mortem digestion. These worms have been found in 13:6 per cent.
of the birds examined, but it must not be forgotten that they
are most inconspicuous and easily overlooked. They have been
found in grouse from Montgomeryshire and Yorkshire, as far
north as Ross-shire, and at all seasons. They do not occur in
large numbers and their pathological effect seems small; still
we must not forget that their near ally, the human parasite
Trichocephalus trichiuwrus (dispar), is one cause of peritonitis
and appendicitis in Man.
However the nematode makes its way into the chick, it must,
like the Zrichostrongylus, grow very rapidly. We have found
specimens in a grouse- chick of fourteen days In age.
(iii.) Family Ascaridee.
(iv.) HETERAKIS PAPILLOSA Bloch.
Stossich * mentions this round-worm, under the synonym of
H. vesicularis Frohl., as occurring in the grouse. It is a very
common parasite in poultry and pheasants. It occurs in the
hand-reared birds at Frimley, and Dr. Cobbett and Dr. Graham-
Smith have found 23 in one cecum and 10 in the other in a
grouse from Abbeystead which was free from 7. pergracilis, and
one in each cecum of a grouse from Longtown, Cumberland, which
had 108 7’. pergracilis in one cecum and 127 in the other, and the
same number similarly distributed in a bird from Bolton Abbey,
which had 3118 7. pergracilis in one cecum and 2877 in the other.
* “Glasnik. Narayosl. druzt.’’ (Societas historico-naturalis Croatica), Zagreb,
1887, p. 284.
1909.} | ‘THREAD-WORMS OF THE RED GROUSE. 349
(iv.) Family Filariide.
(v.) #reartA suiTHT Sambon.
Dr. Sambon * has described a microfilaria or larval form of
some species of Vilaria in the blood of grouse. The adult forms
of such larve usually live in the lymphatics and subcutaneous
tissues; their larvee pass into the blood and are conveyed to new
hosts by some blood-sucking insects.
I am greatly indebted to Dr. E. A. Wilson, net only for
most of the figures which he has kindly drawn, but also for
much help during the working out of the material upon which
this paper 1s based.
EXPLANATION OF THE PLATES.
All the figures have been drawn by Dr. E. A. Wilson. Figures 1 to 27 inclusive
refer to Trichostrongylus pergracilis, figures 28 to 40 inclusive to Tvichosoma
longicolle.
Prate XLVIII.
Fig. 1. Male Trichostrongylus pergracilis, showing alimentary canal, spicules,
and genital bursa. Magnified.
2. Female 7. pergracilis, showing cesophagus, cephalic glands, ovary, uterus
with segmenting eggs, ovejector, and vagina. Magnified.
. Headend of thesame, Showing mouth, oral bulb, pharynx, and cesophagus,
also the cuticular ridges. More highly magnified.
. Tail end of the same. Magnified as fig. 3. Showing anus, rectum, rectal
glands, and posterior loop of uterus.
. Portion of cwsophagus of the same, showing arrangement of cells with
large nuclei forming the cesophageal walls. More highly magnified.
Pratt XLIX.
. Head end of same, showing cephalic glands. Magnified.
. Cuticle of same, one-third from anterior end, showing the ridges. Magnified,
. Vagina and the two ovejectors lying i situ in one straight line, and the
two outer ends of the uteri containing segmenting eggs. Magnified.
. The ovejectors more highly magnified. The thread-worm has been crushed
and the ovejectors have been pressed out and have come to lie parallel
with one another. The lower ends of the uteri with their secreting
eclumnar cells are seen, one of them contains a segmenting egg.
10. Ventral view of posterior end of male, showing arrangement of ribs and of
spicules. Highly magnified.
— oD
Or
Fig.
ie} GO NTS
PuateE L.
Fig. 11. Lateral view of the same, showing the spicules retracted. Highly
magnified,
12. The same, showing the spicules protruded and divaricated. Highly
magnified,
13. The same, seen ventro-laterally, showing attachment of muscles.
14, Various views and stages of spermatozoa.
Puate LI.
Fig. 15. Unsegmented egg.
16. Egg with two blastomeres.
17, Ege with four blastomeres.
18. Egg with eight blastomeres, only six showing.
19. Egg with eight blastomeres, all showing.
20. Ege with sixteen blastomeres.
ea aS AE a oo A SEY ad ES
* Jour. Trop. Med. and Hyg. x. 1907, p. 804. Filaria smithi cannot, however,
stand, as the name is pre-occupied by Cobbold’s species Filaria smithit trom
the Elephant, Tr. Linn. Soc. London, 2nd ser, 11, 1882, p. 237.
ON THREAD-WORMS OF THE RED GROUSE. [ Mar. 16,
Fig. 21. Ege with thirty-two blastomeres.
22, Ege with sixty-four blastomeres.
23. Ege with coiled-up larva, ripe for hatching.
24, A similar egg, artificially ruptured, and the larva in the act of escaping.
Figs.
This shows the contraction of the egg-shell when ruptured.
15-24 inclusive are drawn to the same scale, and the last four or five from
specimens artificially reared in a watch-elass.
Puare LIT.
Fig. 25. A normally hatched embryo, a little older than the stage shown in fig. 23,
found living in droppings of the grouse picked up on a moor.
26. A slightly older larva, possibly of 7. pergracilis, found in the lungs of a
dying grouse.
27, A similar specimen, still in the fiuid of the lungs, but twenty-four hours
after it was last seen alive. This explains why they are so easily
overlooked.
28. Two amceboid parasites in the body-cavity in the neighbourhood of the
ovejector.
29. A female Trichosoma longicolle. Magnified about ten times. (1) Shows
the cesophageal cells, (ii) the prolapsed vagina, (ili) the uterus with eggs,
and (iv) the slightly inflated tail.
Pirate LIMIT.
Fig. 30. Three specimens of same. Magnified about twice.
31. The same, magnified about seven times.
32. Anterior end of the same. More highly magnified.
33. Csophageal cell. More highly magnitied.
34. Uterus with eggs and alimentary canal.
Prats LIV.
Fig. 35. @sophageal cells, pierced by cesophagus, and showing granular nuclei and
to} to} ] y ] to} ) oD
slightly ridged cuticle. Magnified.
36. Optical section, showing right and left glandular bands. Slightly magnified.
37. A glandular band seen im profile; the unicellular glands and their ducts
are dark, the cuticle light. Highly magnified.
38. The prolapsed vagina. Highly magnified. This view shows also the
passage of the cesophagus into the intestine and the neighbouring
glands.
39. Posterior end of the male, showing spicule partially extruded. Highly
magnified.
40. Egg from uterus. Highly magnified.
41. Ege in contents of intestine of the grouse.
PrarE LV.
Fig. 42. Masses of epithelial cells which occur around the 7. longicolle and which
have been broken off the inner walls of the duodenum.
43. The same, isolated and more highly magnified.
44, Nematode larve found, on one occasion, in small intestine of the grouse.
45. The same, more highly magnified.
46. Crystals from rectum.
List of Reference Letters.
a, 6, c. The three chambers of the ove- | o. Ovary.
jector. 0.6. Oral butb.
a.l. Antero-lateral rib. | ce. (Esophagus.
am. Ameeboid, parasitic organisms. oj. Ovejector.
an. Anus. ph. Pharynx.
c.gl. Cephalic glands. p.l. Postero-lateral rib.
d. Dorsal rib. 7. Rectum.
e.d. Externo-dorsal rib. r.C. Rectal cells.
g. Gubernaculum. sp. Spicule.
in. Intestine. | w.&ut. Uterus.
lv. Latero-ventral rib. v. Vagina.
m. Mouth. v.v. Ventro-ventral rib.
ml
. Median-lateral rib.
E.Wilson, Cambridge.
BZ.S.1909. Pl. LVI.
a
-
<j
o
O
fae
5
<
—]
a
<
=
<
=
one D
ae A
IP Avew SOS eit Lill
SS Sy
IW
Sy
: 6
- ~ Se . es <
E Wilson, Cambridge.
DAVAINEA UROGALLI AND HYMENOLEPIS MICROPS.
PAS LOSS Pleven
-B Wilson, Cambridge.
HYMENOLEPIS MICROPS.
7 Se
{
SS SS ee
ow
E Wilson, Gambridge.
DAVAINEA UROGALLI.
PZ.S: 1909) PliLxX
E.Wilson, Cambridge.
DAVAINEA UROGALLI ann HYMENOLEPIS MIGROPS.
LSB iJ ON TAPE-WORMS OF THE RED GROUSE. ool
3. The Tape-Worms (Cestoda) of the Red Grouse (Lagopus
scoticus). By A. H. Surpizy, M.A., Hon.D.Sc., F.R.S.,
F.Z.S., Fellow and Tutor of Christ’s College, Cambridge,
and Reader in Zoology in the University. With a Note
by Wm. Byerave, M.A.
[Received December 18, 1908. }
(Plates LVI.-LX.*)
CESTODA. :
Three species of tape-worm live in the alimentary canal of the
Grouse. The largest of these is (i.) Davainea urogall (Modeer),
which lives in the small intestine (Pl. LVII. fig. 2). We have
also from time to time found it in the ceca; its presence there
is probably due to post-mortem migrations. This is the tape-
worm known to the keepers and to sportsmen generally. It is
large, sometimes a foot or more in length, and is occasionally
seen protruding from the bird’s anus and trailing through the
air as the bird flies. The second and third tape-worms are in-
conspicuous and have hitherto escaped notice. One of them,
(ii.) Davainea cesticillus (Molin), is small and very rare, we have
only found it twice in the many hundreds of Grouse we have
examined. It occurs, a few at a time, in the small intestine.
The third tape-worm, (iii.) Zymenolepis microps (Diesing), 1s also
inconspicuous, and so transparent when alive as almost to be
invisible. It exists in hundreds in the duodenum, and probably
causes a considerable amount of disease and death to the birds
(Pl. LVII. fig. 4). It is by far the most dangerous of the three
tape-worms of the Grouse.
Gi.) Fam. Teeniide.
Genus Davarnua R. BI. & Raill., 1891.
(i.) DAVAINEA URCGALLI (Mcdeer), 1790.
Synonyms: Zenia urogalli Modeer, 1790.
Tenia calva Baird, 1853.
Davainea calva Shipley, 1906,
The worm was apparently named Zwnia wrogalli by Modeer 7
in the year 1790. I am greatly indebted to Dr. O. Fubrmann,
of Neuchatel, for pointing out that this tape-worm is identical
with that described in 1853 by Baird and called by him Tenia
calva.
The following is Baird’s £ description :—
“ Tenia calva Baird, Cat. Entoz. Brit. Mus. 83.
“ Head small, rounded and smooth, white and shining. Mouth
unarmed. Neck constricted. Articulations of body at first very
* Wor explanation of the Plates see p. 362
+ Vet. Ac. Nya Handl. 1790, p. 129.
£ Proc. Zool. Soc. Lond. xxi. 1858, p. 24.
352 DR. A. E, SHIPLEY ON [ Mar, 16,
small, gradually enlarging in breadth as they descend till they
reach about the middle of the body, where they are still narrow,
linear-shaped, and about seven times broader than long. After
this they begin to increase in length and diminish in breadth,
becoming at first nearly square, and at last, near the extremity,
nearly twice as long as broad. All the articulations are strongly
striated across, and the upper and lower margins, where they
join with each other, are considerably thickened. Length
53 inches, greatest breadth 32 lines, breadth of lower extremity
1 millimetre, of head 4 mm.
‘‘ Hab. Intestines of the common Grouse, Lagopus scoticus,
Brit. Mus.”
The same worm has been more fully described, also under the
name 7’. calva, by F. 8. Monticelli*.
The genus Tenia has been comparatively lately broken up into
a number of other genera, and one genus Davainea, named after
the celebrated French helminthologist Davaine, has been estab-
lished for those worms which have the rostellum and suckers armed
with a multitude of characteristically shaped hooks or thorns.
The genus was made in 1891 by R. Blanchard and A. Raiiliet,
and it comprises a number of. species which, as a rule, live in the
smail intestines of birds.
Specimens of Davainea urogalli vary greatly in appearance and
in size. On the whole, they have in life rather an untidy, dis-
hevelled appearance, without clear-cut features ; some preserved
specimens, however, had very definite outlines. Doubtless much
depended on the preservative.
Our longest specimens measured 35 cm. in length ; the greatest
breadth was 4 mm. The preserved material evidently died in
very varying states of contraction, and it is difficult to make
general statements as to the relative proportions of different parts
of the body. One specimen 35 em. in length we found in a bird
of not more than three weeks old. It was shedding ripe pro-
glottides. This worm had split and presented a forked tail,
one limb of which, however, seemed to have dwindled and come
to nothing.
The head is very small. Baird gives its breadth as “1 mm.”
I should put it at about the same, but here, as elsewhere, no two
specimens are exactly alike. The proglottides which follow are
extremely narrow from behind forward, but they very rapidly
increase in breadth, so that 6 or 7 mm. from the head the breadth
is 1 mm., and at about 12 to 15 mm. it is2mm. The greatest
breadth is usually about 2°5 mm. to 3 mm., but in some specimens
4 mm, are reached (Pl. LVII. fig. 2). The broadest portion is
usually about the third quarter of the body from the head ; even
here the segments still have but a very shallow antero-posterior
diameter, about 0-6 mm. to 1 mm. Behind this region the
segments narrow again. They become as long as they are broad,
* Boll. Soc. Napoli, ser. I. v. 1891, p. 155.
1909. | TAPE-WORMS OF THE RED GROUSE. 353
and but for the prominent posterior lip the segments would be
square. The posterior segments are, however, longer than they
are broad, and quite at the hinder end they are attached to one
another by but a slender connection. The prominent posterior
angle is maintained to the last. It is, however, difficult to give
precise statements as to the condition of this worm. In some, one
region of the body will be swollen out; in other specimens, other
yegions will expand. Some have a ‘thin, papery consistence ;
others are plump and almost circular in section. Sometimes
the posterior rim overlaps the anterior region of the succeeding
segment, so that the whole resembles the pile of conical caps which
clowns used—TI do not know whether they still do so—to wear in
the circus. These varying conditions doubtless depend largely
on the state of the parasite when killed and on the means taken
to kill them. The genital pore is, except in rare cases, on one
and the same side.
The anterior end tapers quickly to the very small, squarish head
(Pls. LVI. & LIX. figs.2&6). Anteriorly, the head ends in a ros-
tellum, which seems nearly always to be retracted into a shallow
recess. At each of its four corners the head bears a large sucker,
as a rule circular, but at times oval in shape, and then the long
axis is longitudinal.
Both rostellum and suckers bear hooks, which differ, however,
both in their arrangement and shape. The hooks of the rostellum
are arranged in a double row, Each is shaped something between
a Y and a T (PI. LIX. figs. 7 & 8), one arm being more curved
than the other and it is this arm which is anterior. The stalk
of the hook is but very slightly curved, and the posterior row
alternates with the anterior. The length of the hooks is between
6-9 « and 66. The shape of the hooks does not vary appreciably
and the arrangement in two rings is very regular. ‘These hooks
were not seen “by Baird and were first recognised i fi LEO A,
The hooks on the suckers are also very ‘minute, and they vary
considerably in size: the largest forms are about as long as the
vostellay hooks, 7. ¢. 6°6 p; the smallest forms are perhaps half
this size, and Mena are intermediate sizes. Each hook is slightly
curved and tapers to a fine point, each possesses a “ heel” which,
as is shown in fig. 10, Pl. LIX., is developed in varying degrees.
In many cases tne proximal end resembles the “head” of a
thigh-bone. The hooks are arranged in a ring, but the ring
contains no definite and regularly arranged rows, rather it is a
small circular forest or hedge of hooks of varying sizes and shapes
(Pl. LIX. fig. 9).
The head is usually followed by an unsegmented neck, three or
four millimetres in length, but I have seen one or two specimens
in which the segmentation occurs immediately behind the head.
In transparent specimens longitudinal muscles running to the
suckers can be seen traversing the neck. «In most specimens the
* F.S. Monticelli, Boll. Soc. Napoli, ser. I. v. 1891, p. 155,
354 DR. A. E, SHIPLEY ON [Mar. 16,
reproductive pore is on one side of the body throughout its entire
length, but in others, and rarer, it changes over and having been
for the anterior half of the body on the right side it suddenly
passes to the left and remains there till the end.
The number of the proglottides varies with the length of the
worm. An average-sized specimen would have between 250 and
400 proglottides ; each of these might contain, say, a couple of
hundred eggs. These figures, though necessarily rough, give some
idea of the number of ova a single tape-worm may contain at any
one moment. But mature proglottides are always breaking away
and fresh ones are always being formed, like a recurring decimal,
so that the number of ova a tape-worm produces in the course of
its life is very much greater than the number it contains at any
one moment.
Although the male and female reproductive openings are close
together, the male orifice is very clearly anterior to that of the
female. It leads into a muscular protrusible penis, which was in
all cases retracted. The penis ends in a much coiled vas deferens
which runs half across the proglottis near and slightly obliquely
to the anterior edge ; here it ends in a number of diverticula which
form the testes. These are scattered through the parenchyma.
The wall of the vas deferens is thin, its lumen is spacious, and it
acts as a vesicula seminalis. The lumen is lined by a thin cuticle,
and outside this and all around it are a number of spherical or
oval cells which without exactly forming an epithelium probably
secrete the cuticle.
The vagina opens immediately behind the vas deferens. Its
outermost part is thick-walled and the lumen contains some
homogeneous substance which stains deeply ; further on the wall
becomes thinner, the lumen more capacious, and here masses of
Spermatozoa are to be seen. The ovaries are two, right and left,
each rather of a cauliflower shape; they contain rounded ova in
which besides the nucleus a second deeply staining body is some-
times seen. The vagina makes a turn through about a right
angle, and passing between the two ovaries, where it receives the
two oviducts, it travels back to the vitellarium, a somewhat
pyramidal body lying close to the posterior end of each proglottis.
Certain unicellular glands in this region of the female duct are
probably shell-glands,
There seems to be no walled uterus, but the fertilized ova are
scattered throughout the body embedded in the parenchyma.
Each is a large oval cell with very vacuolated protoplasm and a
nucleus at one side and numerous yolk-granules (Pl. LEX. fig. 12).
Monticelli describes the proglottides as longitudinally striated,
and the striations as due to the longitudinal muscles. These are
certainly conspicuous in section, although in our specimens the
external striation was not very well marked.
One striking feature of D. wrogalli is the great extent to which
the water vascular system is developed. It is spacious and large
in the anterior segments, but in the posterior half of the body it.
1909.] TAPE-WORMS OF THE RED GROUSE. 300
becomes very much larger. The lumen of the lateral canals
increases and the transverse duct which unites them at the
posterior end of each proglottis swells out amazingly. From
being a slender duct it enlarges to a great spherical chamber,
of which the sides, which will rupture when the proglottis drops
off, are extremely thin (Pl. LIX. fig. 13 & Pl. LX. fig. 14).
When the ova are squeezed out of a living ripe proglottis of
D. urogalli, they present the appearance shown in Pl. LIX. fig. 12.
A more highly magnified view is shown in Pl. LX. fig. 15. Hach
ege contains a six-hooked embryo which is much smaller than
the egg-shell. Besides the six-hooked embryo, the egg-shell
contains two or three spherical bodies usually of about the same
diameter as the embryo, but sometimes smaller. These are
apparently yolk-spheres in course of absorption; the remainder
of the ege-shell isempty. The six hooks, arranged in three pairs,
have a characteristic shape shown best in P]. LX. fig. 16. The
shape is similar to that of the hooks figured in the sketches
of Davainea erabryos in Blanchard’s article *
The genus Davainea occurs in many birds, Cursores, Galli-
nace, Columbine, &c., and much more rarely, in the form of
Davainea madagascariensis (Day.), in the intestine ofman. Little
is known of their second hosts; they are usually believed to be
insect larvee, centipedes, or land mollusks. Grassi and Rovelliy
consider the intermediate host of the D. proglottina of the common
fowl to be Limaw cinereus, L. agrestis, and L. variegatus. In this
case the cysticercoid is fully developed in the slug within twenty
days. If theslug be swallowed by a fowl the cy Sticereoid becomes
adult at the end of eight days. We have up till now sought for
the cystic form in Limax flavus without success.
Davainea echinohothridat, which is possibly a synonym of
D, tetragona, causes a nodular disease in poultry §, a condition
liable to be mistaken for tuberculosis. This disease was first
recorded in the United States by Moore (1895) ||, from whose
article the following extracts are made :—
“The nodules were invariably more numerous in the lowest
third of the small intestine. They occasionally appeared,
however, in small numbers in both the duodenum and colon.
The larger and to all appearances older nodules were found in
the ileum near the ceca.
“Tn the badly affected portion the nodules gave the appearance
of closely set protuberances, varying in size from barely per-
ceptible areas of elevation to bodies 4mm. (+ inch) in diameter.
In some instances they appeared to overlap oneanother. When
separated by a band of normal tissue they were round or some-
what lenticular in form. In the latter case the long diameter
* Mém. Soc. Zool. France, iv. 1891, p. 420.
+ Centrbl. Bakter. iii. 1888, p. 172.
+ B. H. Ransom, ‘Manson’s Eye Worm of Chicken, &c.,’ Bureau of Animal
Industry, U.S.A., Bulletin 60, 1904.
§ D. HE. Salmon, y Tapeworms of Poultry,’ Bureau of Animal Industry, U.S.A.,
Bulletin 12, 1896.
Ie WieeAs Moore, Bureau of Animal Industry, U.S.A., Circular I1I., 1895.
356 DR. A. E, SHIPLEY ON [ Mar. 16,
was usually transverse to the long axis of the intestine. The
larger nodules were of a pale dark- yellowish colour, while the
smaller ones varied in shade from the more highly coloured
areas to the neutral grey of the normal serosa, ‘lo the touch
they gave the sensation that would be expected if the subserous
and muscular coats were closely studded with small, oval; solid
bodies. ‘The mucosa presented similar elevations. Attached
to the mucosa over the nodules were a number of tapeworms.
There were also in the more advanced cases a variable number
of small (0°5 to 1 mm.) areas over the larger nodules in which
the mucosa had sloughed, leaving small ulcerated depressions.
“The larger nodules contained a greenish-yellow necrotic
substance, Which appeared in the advanced stages as a seques-
trum with a roughened surface. On section it has a
glistening, homogeneous appearance. Surrounding the ne-
crotic substance was a thin layer of infiltrated tissue. The
smaller nodules contained a more purulent-like substance and
the smallest appeared to the naked eye as areas of infiltration.
Sections of the affected intestine showed upon microscopic
examination that the heads of the tape-worms had penetrated
the mucous membrane and were situated in different layers of
the intestinal wall (cf Pl. LX. fig. 17). They were frequently
observed between villi. As would be expected, the heads were
not readily detected in the necrotic masses contained in the
larger nodules, but were almost invariably seen in the smaller
ones. Ina few sections the tape-worm could be traced through
the mucosa to the nodule in the muscular tissue in which its
head appeared, In the earlier stage of the nodular development
there is a cell infiltration about the head of the worm, This
process continues until the infiltrated tissue reaches a con-
siderable size.
* The worms attached to the mucosa were usually small. A
larger form was commonly found in the intestinal contents.
Although macroscopically they appeared to be different,
Doctor Stiles found that they were presumably of the same
species.
* Keonomie Importance.—The importance of this disease is
much greater than it at first appears, as the close resemblance
of the nodules to those of tuberculosis renders it of much
significance from a differential standpoint. As the intestines
are stated to be frequently the seat of the specific lesions of
tuberculosis in fowls, it is of the greatest importance that a
thorough examination be made before a positive diagnosis is
pronounced. There are already several statements concerning
the presence of tuberculosis in fowls in which the data given
are not suttiicient to differentiate the disease from the one here
described. A somewhat analogous disease of sheep caused by
a nematode (Z’sophagostoma columbianum Curtice) has led to
the deliberate destruction of many animals, the owners believing
that tuberculosis was being eliminated from their flocks,
1909.) TAPE-WOBRMS OF THE RED GROUSE. 357
“ As the inquiry into the cause of poultry diseases becomes
more general it is probable that this affection will be occasion-
ally encountered, and unless its nature is recognized it may in
some instances, like the sheep disease, lead to an unwarranted
destruction of property.
“ Tn addition to its importance in differentiating tuberculosis
it is in itself a malady worthy of careful attention. The fact that
- it has already appeared in two flocks in the Distriet of Columbia,
and also in the States of North Carolina and Virginia, shows
that the infesting cestode is quite widely distribute] in this
eountry. It is highly probable that the total loss it occasions,
both from deaths and from the shrinkage of poultry products,
due to the chronic course of the disease it produces, is very
large.”
(ii.) DAVAINEA CEesTICILLUS (Molin), 1858.
Synonym: Tenia cesticillus (Molin).
This is a small species; the majority of our specimens measured
between 5°5 and 9 mm. in length. Few were longer, though many
were shorter. They were all young immature specimens. The
broadest at their broadest part, usually about the level of the last
proglottis but two or three, measured 1 mm. across. They tapered
to the last proglottis, which averaged about 0°5 mm. in diameter,
and still more do they taper towards the head, where the narrow
neck is but 0°2 mm. The head itself is 0°3 to 0°5 mm. across and
perhaps two-thirds of this in length.
The hooks in the rostellum were numerous, I should judge a
few hundred, but I could not, on account of their minute size,
count accurately: they measured about 7 p in length.
The head when the rostellum is withdrawn is somewhat cup-
shaped and the four suckers are on the edge of the cup, opening
at the edge and slightly inwards. There is practically no “ neck,”
just a constriction between the head and the first proglottis.
Behind the head the proglottides increase markedly in size, and
the third proglottis in most specimens is already as broad as the
head. They are deeply imbricated and the overlapping edge is
full and rounded. At the level of the anterior end of each pro-
glottis is a constriction which slightly separates off the overlapping
lobe from the preceding proglottis, to which it, of course, belongs.
This gives a somewhat ear-like outline to the side of each seg-
ment. The constriction first appeags in about the tenth proglottis,
and the characteristic outline is lost in the last, where the over-
lapping edges curve in asif to guard the excretory opening. The
total number of the proglottides varies a little with the variable
length, but differences in length depended far more on the state of
contraction of the body than on the number of segments.
Roughly speaking, the numbers varied from about eighteen to
about twenty-eight proglottides.
308 DR. A, E, SHIPLEY ON [ Mar. 16,
The genital pore is alternate and fairly regularly so; the penis
often projects, and then it is apparent that the pore lies rather
anteriorly andis all but overlapped by the imbricated edges of the
proglottis next in front.
Sections show that there are a number of calcareous bodies in
the tissues; some of these are in optical section brick-shaped,
and others spherical or shaped like a cottage-leaf. These latter
are bigger than the others and show numerous radiating lines.
Posteriorly the tissue becomes very highly vacuolated and the
embryos lie in small packets which do not seem to be in a uterus,
and may be, as Morell suggests in J. wrogalli, inthe lumen of the
ovary itself.
This tape-worm, common in chickens and turkeys, is only an
occasional parasite of the grouse, and has in many hundreds of
birds we have examined only been found twice, and in neither
case has its presence been associated with any lesions. As a
factor in grouse-disease it may be neglected. in both cases only
young, immature, not fully grown specimens were met with.
Its second host is according to Railliet, quoting Grassi and
Royelli, probably some Coleopteran or Lepidopteran, but at present
this has not been proved.
Hymencieris Weinland, 1858,
(ii.) Hymenotepis microps (Diesing), 1850.
Synonyms: Zenia microps (Diesing), 1850.
Hymenolepis tetraonis (Wolfth.), 1900 *.
This is an extremely delicate, transparent tape-worm which
exists in almost countless numbers in the duodenum of Lagopus
scoticus. It is also recorded from the Blackcock and the Caper-
caillie. On cutting open the duodenum of a grouse infested with
these worms—and we have rarely found a bird free from them
except in the winter months—they are not at first apparent.
They are so fine and so transparent that they are practically
invisible when alive, and the contents of this part of the
alimentary canal appears very much like a thick purée. If we
add to this some fixing agent such as corrosive sublimate this
purée resolves itself into a mass of very fine, delicate, white
threads inextricably tangled up together and so numerous that
there seems but little room left in the duodenum for the passage
of the food (Pl. LVIITI. fig. 3). Tf, with great care—for they break
at the slightest strain—we succeed in disentangling one of these
worms we shall find its head embedded to a greater or less extent
in the mucous lining of the duodenum, into which, to use a
poetic phrase, “it nuzzles” whilst the body of the worm floats
freely in the fluid contents of this part of the alimentary canal.
Tf we also succeed in freeing the head we now have a complete.
worm and can study its structure.
* Wolffhiigel, K., “Beitrag zur Kenntniss der Vogelhelminthen,” Inaug.-Diss.,
Freiburg-1.-B. 2900.
1909. | TAPE-WORMS OF THE RED GROUSE. 359
Before giving some anatomical details of H. microps it is worth
mentioning that Wolffhiigel found fragments of this species—
none with the head—in the small intestine, large intestine, and
end—he does not say which end—of the ceca of Tetrao urogallus.
We have also found short chains of ripe proglottides passing
down the alimentary canal on their way to the exterior, but the
tape-worm as an individual lives only in the duodenum,
H, microps is a very long worm, attaining in the longest
examples a length of some 15-16 ems. It consists of an enormous
number of proglottides. The first two millimetres which come
after the head contain as many as 60-70 segments, and lower
down the body, where the proglottides were mature, as many as
10 proglottides measured but 1 mm. Of course these measure-
ments depend entirely on the state of the contraction of the
worm, but if we take the mean between them as a rough average
approximation we shall get the astonishing number of 3000
proglottides in a single specimen. As each proglottis contains a
large number of eggs and as they are being continually renewed,
and as, further, the number of tape-worms in the duodenum
amounts to hundreds, it is easy to see that a grouse-moor must
be just peppered over with ova (Pl. LVILI. fig. 4).
The head is somewhat squarish (Pl, LX. fig. 18), with a central
retractile rostellum and four suckers at the corners. The ros-
tellum is surrounded by a closely packed ring of very numerous
spines or hooks (Pl. LX. figs. 19 & 20). These are very minute
and, except in the fresh specimen, very ditticult to see, and even
then it requires an immersion-lens to make out anything of their
structure. Their proximal end is rounded, and then comes a
constriction ; the spine then thickens till about the middle of its
length and then tapers to a very fine point. Although these
spines are slightly curved, they are in no sense hooked (PI. LX.
fig. 20). I have tried to measure the length of these spines from
specimens of the head, which has been cut in sections. I am not
quite sure that the hooks were entire, and so am not quife sure
that my measurement is large enough, but I should put their
length at avout 16 ~—certainly not less. The hooks seem to be in
a single row, but very close together.
The suckers are deep and well marked, but it must always be
borne in mind how very small the head is, and corresponding
with this the suckers are also very minute.
The posterior edge of each proglottis is “ sailliant,” but it does
not overhang the succeeding proglottis; it stands out like the
tooth of a saw, and viewed laterally the side of this worm is very
saw-like. Throughout the body the proglottides are much
broader than they arelong. In the older ones there are numerous
calcareous bodies, the measurements of which Wolffhiigel gives as
0:018 mm. by 0°01 mm. (Pl. LX. fig. 21).
The genital pore is in all the segments on the same side; the
left, judging by the orientation suggested by the female repro-
ductive organs, being on the ventral surface. The vagina opens
360 DR. A. E. SHIPLEY ON [ Mar. 16,
into a peculiarly large and muscular receptaculum seminis, which
runs across the proglottis and then turns backward; in some
preparations this turn is seen “en face” and then the radiating
muscles give the appearance of a ring of very fine spines, and,
indeed, at first I thought that there was such a ring, but I believe
the above is the true explanation. ‘There are three testes the vasa
deferentia of which unite and after entering the cirrus-bulb enlarge
to form a vesicula seminalis. The vagina opens ventral to the
penis. The uterus isa single chamber unbranched. It forms a
conspicuous feature in the hinder end of stained specimens. At
first it appears as a spherical organ lying in the middle line at
the hinder end of each proglottis, but as it grows and absorbs
more of the parenchyma it tends to become triangular or square,
but always with very rounded angles. It contains a large number
of relatively large onchospheres or tape-worm embryos (PI. LX.
fig. 22). According to Wolfthiigel, the embryos measure 0-02 mm.
in breadth by 0°04 mm. in length. The typical six embryonic
hooks are very characteristic. The partners in each pair, for
instance, are usuaily widely divaricated ; their length is 0-014 mm.
These characteristic Hymenolepis ova have three envelopes: the
innermost, closely applied to the embryo, is never produced into
horns; between it and the middle envelope is only a clear fluid in
which the embryo floats; between the middle and the outer
envelope are the much vacuolated remains of cells. The position
of the embryo is eccentric with regard to this outer shell (Pl. LX.
fig. 22), which measures 0-073 mm. by 0-066 mm. The measure-
ments are again Wolfthiigel’s. The characteristic hooks are
figured on Pl. LX. fig. 23.
We have no information about the fate of these embryos,
but as a general rule the cystic form of this genus lives in some
Insect or Myriapod, as is shown by the fact that this genus of
tape-worm occurs in Bats, Insectivores, Rodents, and Insectivorous
birds. Hymenolepis nana occurs in man, most frequently in
children, and is not at all uncommon in Italy. Sporadic eases of
H, diminuta occurring in man are also recorded.
We have made and we are making laborious investigations to
try and discover this second host. In searching for the cysts
of the Tape-worms we began with the insects which occurred
most commonly in the crop of the grouse. These we examined
microscopically, both after teasing the body up in glycerine and
by grinding it up—but not too finely—in a pestle ; in some cases
also, as Mr. Fryer* has recorded, sections were made and
examined, but always without result.
We were at two disadvantages in hunting for the cysts:
firstly, we did not know what the cysts of either Davainea
urogalli or Hymenolepis microps were like; and,secondly, the tissues
* Interim Report of the Grouse Disease Inquiry.
1909. ] TAPE-WORMS OF THE RED GROUSE. 361
of the insects and spiders which we examined are little, if at all,
known, and more than once we have at first sight taken some
organ proper to the insect for a cestode cyst, only to our great
disappointment to discover later that we were looking at an ovum
or other structure belonging to the putative host.
During some days Dr. Wilson and I spent in Edinburgh towards
the end of July, 1908, we examined a considerable number of the
commoner insects found on the moors in the hope of throwing
some light upon the life-history of the tape-worms so common in
the grouse. The specimens we investigated were collected by
Mr, P. H. Grimshaw, who is preparing a Report on the Insects of
the Moors. We are greatly indebted to him and to the Keeper
of the Museum, Mr. W. Eagle Clarke, and to Mr. J. Ritchie for
kindly placing at our disposal a work-room and other accommoda-
tion which greatly facilitated our work. When the insect had not
been specifically named we always kept a similar specimen for
subsequent identification in case it should contain the cyst; but,
alas ! here again our jabour was in vain.
In the manner indicated we examined the following Insects, in
every case looking through the débris of some four or five
specimens.
DIprera.
(i.) Monophilus ater, one of the subfamily Limnobiinz of the
Tipulide. A very common constituent of the food of young
grouse. No trace of a cyst was found, but in one specimen an
immature nematode was wriggling about.
(ii.) Bibio sp. Here again we drew a blank.
(iii.) Cyrtoma spuria, one of the Empidee. This fly is small and
seemed to have little interior ; no trace of a cyst was found. In
another small Empid fly we discovered a Gregarine.
(iv.) Scatophaga sp. Scatophaga stercoraria is perhaps the
commonest fly in Scotland, and, owing to the larva living in the
droppings of the grouse, it can hardly fail to contain the egys of
the cestodes ; but we have never found a Scatophaga in the crop
of a grouse, and there is some reason to doubt if the tape-worm
eggs develop in this fly. After searching for a long time
through the tissues of many specimens of Scatophaga, we only
managed to find one ovum, apparently of Davainea wrogalli, and
that was no further advanced than when it was laid.
PLECOPTERA.
Similar gropings through the disjected membranes of an un-
known species of Perlid produced no better results.
ARACHNIDA,
We also investigated the tissues of a spider very common on
the moors, and of a phalangid, with an equal want of success.
Proc. Zoou. Soc.—1909, No. XXIV. 24
362 ON TAPE-WORMS OF THE RED GROUSE. | Mar. 16,
Nore sy Wm. Byarave, M.A., oN THE SEARCH FOR Cysts.
Since September 1908 I have been making a series of investi-
gations in connexion with the Grouse-Disease Inquiry. My work
has consisted of a careful examination of the tissues of certain
insects found on Grouse-moors in various parts of England and
Scotland, the object being to discover, if possible, cysts of the three
species of tapeworm which infest the grouse, viz. :—
Davainea wrogalli (Modeer, 1790).
Davainea cesticillus (Molin, 1858).
Hymenolepis microps (Diesing, 1850).
The insects examined to date are specimens of Scatophaga
squalida from Ballindalloch, and S. stercoraria from Burley,
Dunachton, and Forrigen.
- The specimens were sent to me by Mr. P. H. Grimshaw, from
the Royal Scottish Museum, Edinburgh, preserved in spirit.
The method of examination was as follows :—
The legs and head were removed and the body of the insect
teased up in 70 per cent. alcohol as finely as possible with needles,
the legs and head being firstly teased and then gently pounded in
a mortar.
The material thus obtained was examined under a cover-glass,
a mechanical stage being used to ensure that none of the material
was overlooked. The powers used were Leitz Obj. 1” and 3”
Oc. 2 and 4; an oil immersion-lens being used in cases of doubt.
So far the examination has yielded no results. Nothing has been
found which in any way resembled the cysts, one or two of
which have been figured, of species allied to the three tape-worms
mentioned above.
EXPLANATION OF THE PLATES.
Pratt LVI.
Fig. 1. View of a portion of the small intestine of the grouse well infested with
Davainea wrogalli (Modeer) and cut open to show the worms.
Pratre LVII.
Fig. 2. A single specimen of D. wrogalli, isolated to show the attenuated head.
4. A single specimen of Hymenolepis microps, isolated.
(The figures 2 and 4. are very slightly magnified.)
Prare LYIII.
Fig. 3. View of a portion of the duodenum of the grouse, well infested with
Hymenolepis microps (Diesing), cut open to show the worms.
Prats LIX.
Fig. 5. Head of D. wrogalli with proboscis half-exserted and armed suckers,
6. The same, with the proboscis completely retracted. This specimen shows
the calcareous bodies well.
7. Portion of the double circle of hooks from the proboscis of D. wrogalli.
8. Isolated hooks from the same, showing slight modification in outlines.
9. Portion of the ring of hooks which surrounds one of the suckers of
D. urogalli, showing the irregular arrangement of the hooks.
10. Isolated hooks from the same, showing slight modification in outline.
1909.] ON PARASITES OF BIRDS ALLIED TO THE GROUSE. 363
Fig. 11. Magnified view of mature proglottis of D. wrogalli. gp., genital pore;
_-0v., OVary: p., penis; ¢., testes; v., vagina: vié., vitellarium ; w.v., excre-
tory system. ;
12. Ova of D. urogalli, showing the ovum and yolk-vesicles.
13. Longitudinal horizontal section through the same, showing the great
extension of the excretory transverse canals, and the ova scattered zn egg-
capsules.
PratEe LX.
Fig. 14, A similar section through the last proglottis, showing the enormous trans-
verse canal at the level of the breaking zone and the excretory pore e.p.
15. Ova of D. urogalli more advanced than those shown in fig. 12, showing the
onchosphere and traces of the yolk-vesicles.
16. An isolated hook from the same.
17. Transverse section of the walls of intestine of a fowl, showing Tenia
botriopliti embedded in the deeper layers of the intestinal wall. aa., in-
testinal mucosa; 6., muscular layers; sp., peritoneal lining; ¢¢., anterior
ends of the tape-worms; ¢., mass of exudate produced by the irritation of
the head of the Tenia. (From Piana, Mem. Ac. Sci. Istit. Bologna, series
4, vol. 11. 1880, p. 387.)
18. Head of Hymenolepis microps (Diesing), highly magnified.
19. Section through the retracted proboscis of the same, highly magnified to
show the arrangement of the hooks.
20. Isolated hooks of the same seen under {5 oil immersion-lens.
21. A few mature proglottides of the same, taken from about the middle of the
body.
22. Onchospheres of H. microps in the characteristic three envelopes.
23. Hooks from the same.
4, Internal Parasites of Birds allied to the Grouse. By
A\. Jd, (Suni, Wiel deloms IDS, IeltSn Jena,
Fellow and Tutor of Christ’s College, Cambridge, and
Reader in Zoology in the University.
| Received December 18, 1908. |
The following is a brief enumeration of the Cestode, Trematode,
and Nematode parasites of the Grouse, the Ptarmigan, the
Blackeock, and the Capercaillie. To these I have added the
Willow-grouse and the Hazel-hen, although these birds, unlike
the three former, are not denizens of the British Isles. It will be
noticed that but two Nematodes, 7richosoma longicolle (Rud.) and
Heterakis papillosa (Bloch), and two Cestodes, Davainea wrogalli
and Hymenolepis microps, are common to our grouse and to its
allies. The first named round worm and the tape-worm are
found in all three, Blackcock, Capercaillie, and Grouse, and in none
of the other nearly allied birds; whilst Heterakis papillosa has
been recorded from the Ptarmigan, the Capercaillie, the Hazel-
hen and the Grouse, besides from many other birds.
Heterakis perspicillum (Rud.) also occurs in three hosts, in the
Blackeock, the Hazel-hen, and the Capercaillie. All the other
Nematode parasites mentioned occur in a single host, except
Heterakis compar (Schrank) which is found in both the Ptarmigan
and the Capercaillie. Of the Cestodes, Davainea wrogalli and
Hymenolepis microps occur in the Grouse, the Blackgame, and the
Capercailhie.
24%
364 DR. A. E, SHIPLEY ON INTERNAL PARASITES — [| Mar. 16,
I. Lacorpus mutus Leach, or L. auprnus. The Ptarmigan.
The Ptarmigan contains Heterakis (Ascaris) compar (Schrank),
Heterakis papillosa (Bioch), and Heterakis borealis (v. Linstow),
amongst the Nematodes, and Tenia echinatw amongst the
Cestodes.
NEMATODA.
(i) Heterakis compar (Schrank).*
Synonyms. Ascaris compar Schrank 1790.
Ascaris lagopodis Frohlich 1802.
Fusaria compar Zeder 1803.
Heterakis compar Stossich 1888.
A thin, white Nematode, the females of which attain a length
of 84-96 mm., and the males a length of 36-48 mm., found
in small intestine of the Ptarmigan, the Capercaillie, and other
gallinaceous birds.
(ii) Heterakis papillosa (Bloch).
Synonyms. Ascaris papillosa Bloch 1872, nec Molin 1860.
Ascaris vesicularis Frohlich 1791 pro parte.
Heterakis vesicularis Duj. 1845.
Heterakis papillosa Railliet 1885.
The male measures 7-13 mm., the female 10-15 mm. <A very
common parasite of the ceca of the alimentary canal. Some-
times it exists in prodigious numbers, causing a fatal perityphlitis
amongst chickens.
This parasite is common in domestic fowls, ducks, geese, turkeys,
peacocks, guinea-fowls, pheasants, etc., and occurs in the
Ptarmigan, the Capereaillie, and the Hazel-hen (Lonasia sylvestris).
The eggs of H. papillosa complete their development in water.
(iii) Heterakis borealis (v. Linstow).
Very little is known about this nematode, which was described
in 1884 by von Linstow in the ‘Archiv fiir Naturgeschichte,’
Nee Tos Laalle
CESTOD A.
(i) Tenia echinata (Olss.).7
Il. Lacorus ausus, or L. SUBALPINUS, or L, sALicETI. The
Willow-grouse. I can find no record of any parasites being
recorded from the Willow-grouse.
* Where possible the nomenclature follows that of Railliet in his “ Zoologie
Médicale et Agricole.”
+ Olsson, “ Bidrag til Skandinav. Helminthfauna,” ii., Stockholm, 1893.
1909. ] OF BIRDS ALLIED TO THE GROUSE. 369
III. Bowasta sytvestris. The Hazel-hen.
Tetrao bonasia Linn.
NEMATODA.,
(1) Heterakis papillosa (Bloch).
v. under Lagopus mutus, p. 364.
(ii) Heterakis perspicillum (Rud.).
Synonyms. Ascaris gallopavonis Gmelin 1789.
Fusaria refleca Leder 1800 pro parte.
Fusaria strumosa Geder 1800.
Ascaris perspicillum Rud. 1803.
Ascaris gihbosa Rud. 1809.
Ascaris infleca Rud. 1819 pro parte.
Ascaris funiculus KE. Deslongchamps 1824.
Heterakis inflecu Schneider 1866.
Under the last synonym this worm has recently been recorded
from the alimentary canal of a Russian Hazel-hen by Wolffhiigel*.
The males measure 3-8 ems., the females 6-12 ems. The worm is
common in fowls, guinea-fowls, and turkeys, and gives rise to
severe epizootics amongst poultry. Occasionally they are found
free in the body- cavity, and sometimes occur in eggs, having
made their way into the ovum before the shell is deposited.
(i) Milaria bonasie (v. Nordmann).
This form apparently has been seen but once. It was found by
v. Nordmann in the posterior chamber of the damaged eyes of a
Hazel-hen taken in the island Wikari.
CESTODA.
Ga) Tenia bonasie (Miller).
This tape-worm is recorded from the intestines, but practically
nothing is known about it.
TV. Terrao terrix L. Blackeock.
NEMATODA.
G) Trichosoma longicolle (Rud.).
This is the worm (or one of them, more than one species may
be included in Rudolphi’s original description) described by
Rudolphi. It occurs in the fowl, pheasant, and grouse as well as
in blackgame, v. p. 345.
(ii) Heterakis compar (Schrank).
v. p. 364.
* “ Beitrag zur Kenntniss der Vogelhelminthen,” Inaug.-Diss., Freiburg i. B., 1900.
+ Miller, Arch. Naturg. 1897, p. 10, and Stossich, “‘ Glasnik. Naravos]. druzt.”
1887, p. 287.
366 DR. A. E, SHIPLEY ON PARASITES [ Mar. 16,
Gili) Heterakis perspicillum (Rud.).*
v. p. 365.
(iv) Heterakis magnopapilla (v. Linstow). T
(v) Strongylus papillatus (v. Linstow). £
Wolfthigel found both males and females in great numbers in
the alimentary canal of the Blackeock. It had previously been
described by v. Linstow from the intestine of the Great Bustard,
Otis tarda.
(vi) Strongylus hastatus (v. Linstow). $
(vii) Nematode % genus, recorded by Wolffhiigel from the
ceca.
TREMATODA.
(1) Lyperosomum corrigia (Braun). |)
A fluke nearly allied to Distomum plesiostomum (vy. Linstow,
1883) from Perdix greca.
CESTODA.
(i) Davainea retusa (Clerc). &|
Gi) Davainea urogalli (Modeer).
Described by Krabbe from specimens in Perdix greca, Megalo-
perdix nigelli, Tetrao urogallus and T. tetria.
(i) Davainea villosa (Bloch).
Also found in a Megaloperdix nigelli in Turkestan.
(iv) Hymenolepis microps (Diesing).**
This species, described by Wolffhiigel under the name of
H. tetraonis from a few fragments, is more fully described by me
from Lagopus scoticus in the anatomical part of this memoir,
cf. p. 358.
A CANTHOCEPHALA.
(i) Echinorhynchus stellaris (Molin).
Usually found in the Duck, but Dr. von Linstow tells me he has
seen it in the larval stage encysted on the outside wall of the
intestine in the Blackcock.
* “Glasnik. Naravosl. druzt.” (Societas historico-naturalis Croatica), Zagreb,
1887, p. 278.
+ “Schr. Ges.” Konigsb. xlvii. 1906, p. 112.
Arch. Naturge, 1882 (i), p. 3.
Tbid., 1905, p. 274.
|| Centrbl. Bakter. xxix. 1901, p. 946.
@ Rev. Suisse Zool., xi. 1903, p. 363.
=* “ Beitrag zur Kenntniss der Vogelhelminthen,” Freiburg i. B., 1900.
1909.] -
OF BIRDS ALLIED TO THE. GROUSE.
367
V. Terrao uvrocautus L. The Capercaillie.
WNEMATODA.
(i) Heterakis compar (Schrank), v. p. 364.
(ii) Heterakis papillosa (Bloch), v. p. 364.
(ii) Heterakis perspicillum (Rud.), v. p. 365.
(iv) Filaria urogalli (v. Linstow).
Lives in the subcutaneous tissues.
(v) Trichosoma longicolle (Rud.), v. p. 345.
CESTODA.
(1) Bothriocephalus sp.
Cobbold* found in the subeutaneous tissue lying over the
pectoral muscles of a Capercaillie a single specimen of a tape-worm
which he compared with a Ligula reptans.
larval Bothriocephalus.
It was probably a
(ii) Davainea urogalli (Modeer), v. p. 351.
(ui) Davainea globocaudata (Cohn).
(iv) Hymenolepis microps (Diesing).
List oF PARASITES WITH THEIR Hosts.
NEMATODA.
Filaria bonasie (v. Nordm.) ...
Filaria wrogalli (v. Lins.)
Filaria sp.
FHeterakis borealis (v. Lins.)
Heterakis compar (Schrank) ...
Heterakis papillosa (Bloch)......
Heterakis perspicillum (Rud.)...
Feterakis magnopapilla (v. Lins.)
Strongylus papillatus (v. Lins.)
Strongylus hastatus (v. Lins.)...
Syngamus trachealis (v. Sieb.)
Trichosoma longicolle (Rud.) ...
JHOS I,
Bonasia sylvestris.
Tetrao urogallus.
Lagopus scoticus.
Lagopus mutus.
Lagopus mutus, Tetrao tetrix,
and Tetrao wrogallus.
Lagopus mutus, Lagopus scoti-
cus, Tetrao wrogallus, and
Bonasia sylvestris.
Tetrao tetrix, Tetrao urogallus,
and Bonasia sylvestris.
Tetrao tetria.
Tetrao tetria.
Tetrao tetria.
Lagopus scoticus.
Tetrao tetrix, Tetrao wrogallus,
and Lagopus scoticus.
Trichostrongylus pergracilis (Cobb.) Lagopus scoticus.
Nematode gent | Ae eeteseceee
Tetrao tetrix.
* Tr. Linn. Soc. London, xxii. p. 165.
7+ Zool. Anz, xxiii. 1900, p. 91, and Acta Ac. German. Ixxix. 1901, p. 263.
368 MR, W. P. PYCRAFT ON A [ Mar. 16,
TREMATODA. HOST:
Lyperosomum corrigia (Braun) Tetrao tetrix.
CESTODA.
Bothriocephalus sp. 0.1... +1200 Tetrao urogallus.
Davainea cesticillus (Molin) ... Lagopus scoticus.
Davainea globocaudatus (Cohn) Tetrao wrogallus.
Davainea retusa (Clere) ......... Tetrao tetrix.
Davainea urogalli(Modeer) ... Tetrao tetrix, Tetrao urogallus,
and Lagopus scoticus.
Davainea villosa (Bloch) ...... Tetrao tetrix.
Hymenolepis microps (Diesing) Tetrao tetrix, Tetrao urogallus,
and Lagopus scoticus.
Tenia bonasie (Miull.) ......... Bonasia sylvestris.
Teenia echinata (Olss.) ......... Lagopus mutus.
ACANTHOCEPHALA.
Echinorhynchus stellaris (Molin) Tetrao tetria.
5. On a Fossil Bird from the Lower Pliocene.
Bay Weve MORARM INH: Zse, cA ease
[Received February 16, 1909. ]
(Text-figure 47.)
The following aceount concerns the fossilized remains of a
small Passerine bird from the Lower Pliocene of Gabbro, near
Leghorn. The slab in which these remains are embedded was
placed in my hands for investigation by my friend Dr. Forsyth
Major, F.R.S., who has, throughout my enquiry, rendered me
much help.
Unfortunately, only the pelvic limbs and a few traces of
feathers are here preserved ; further, the bones are much crushed,
and the phalanges have been almost entirely lost. But from the
slab and its counterpart, which has happily been preserved,
sufficient details may be gathered to make identification possible.
The right leg lacks the toes. The femur, on the slab, is much
crushed and can only very imperfectly be traced; but on the
counterpart of the slab it becomes clear that it is seen from its
dorsal aspect, since the middle of the shaft shows a smooth,
periosteal, surface; while the extremities thereof are missing,
leaving beautiful impressions of the distal and proximal articular
ends. The tibio-tarsus is seen from its fibular side. Herein traces
are visible of the head of the fibula, closely approximated to the
femoral trochlea, and of the external border of the ectocnemial
* Published by permission of the Trustees of the British Museum.
1909. | FOSSIL BIRD FROM ITALY. 369
crest. The fibular crest has been crushed. ‘The distal end of the
shaft, though crushed, shows the form of the trochlea. On the
counterpart the proximal extremity of the shaft is much damaged ;
while of the distal end only an impression remains.
Text-fig. 47.
The Slab (left-hand figure) and the counterpart thereof (right-hand figure)
containing remains of a fossil Pipit (Anthus bosniaskii).
The hind-toe of the left leg is clearly seen in the Slab, while to the left of the tibial
shaft of the right leg traces of feathers are preserved (marked x), but these are
much more clearly seen on the counterpart.
The left leg is more complete, but lacks the front toes. On
the slab the femur is seen from its ventral aspect; the head
is missing, but the condyles of the distal extremity can be fairly
well made out. The tibio-tarsus is seen from its inner aspect.
It is therefore somewhat curious to find that the fibula has become
detached from the fibuiar crest, and les with its head in close juxta-
position to the fibular condyle of the femur, as may be seen in
the photograph (text-fig. 47). The entocnemial crest is fairly well
preserved. The counterpart of this slab lacks the proximal ex-
tremity of the tibio-tarsus, owing to the flaking away of the matrix,
while the distal end of the shaft can only very indistinctly be
made out. The tarso-metatarsus, though preserved in its entirety,
is too much crushed at its proximal extremity to make it possible
to distinguish the mesotarsal articulation. The shaft is also
flattened and crushed. The ecto- and entocnemial trochlez can
be made out, but the mesotrochlea is obliterated.
Of the digits only the hallux is preserved, but the metatarsal
thereof cannot be traced, while the ungual phalanx is much
erushed and flattened. This digit, however, serves at once to
show the Passerine character of these remains, and this because
of the great length of the proximal phalanx and its position in
regard to the other toes. Of the other digits only fragments of
370 ON A FOSSIL BIRD FROM ITALY. [Mar. 16,
the proximal phalanges remain. The only trace of the tarso-
metatarsus and hallux, which is preserved in the counterpart,
takes the form of a very shallow depression in the matrix,
showing that an overlying flake of stone and the bone imbedded
therein have been lost.
There is a cluster of fragments of other bones in this slab, and
these appear, at first sight, to be portions of thoracic vertebree—
crushed centra and neural spines. A closer examination shows that
these “spmes” are too high to belong to the vertebral column
of any Passerine bird of this size.
And now a word as to the traces of feathers which are to be found
on the slab and counterpart containing the bones just described.
These impressions occur in the form of a large patch occupying
the middle of the area shut in by the femur and the shank of
the left leg. On the slab a portion of this patch has been lost,
but in the counterpart it is complete, and indicates a mass of
feathers, probably of the flanks, matted together as if by wet.
They show, moreover, that these remains must have been exposed
to the action of tides, where low water would leave the feathers
in a matted “draggled” condition. Before finally covered by
the next high tide they must have become effectually covered
by mud ; antl this because feathers under water would float out,
much as in life.
Along the lower end of the “shank” of each leg there are
also impressions of feathers, which cease at the mesotarsal
articulation.
SUMMARY.
There seems little room for doubt but that the remains just
described are those of a Pipit (Anthus); at any rate, they agree
more closely with bones of this genus than with those of any
other group of Passerine birds with which I have compared
them. And among the members of this genus these remains
resemble most nearly those of the living species known as
Berthelot’s Pipit (Anthus bertheloti).
In the shape of the condyles of the distal end of the femur,
and of the trochlez of the distal extremity of the tarso-metatarsus,
shght differences are discernible when the fossil remains and
the skeletons used for the purposes of comparison in this paper
are compared. But these differences are very slight, and they
may be due to pressure.
Assuming that I am correct in ascribing these remains to an
extinct species of the genus dnthus, 1 propose to adopt the name
of the discoverer—Dr. Bosniaski—as the specific name. <Anthus
bosniaski was obtained by Dr. Bosniaski from the Lower Pliocene
of Gabbro near Leghorn, a deposit which has yielded many fossils
and which is particularly rich in fish-remains.
So far as I can make out, the only other remains of Passerine
birds from the Lower Phocene are a few fragments representing
the genera Corvus and Turdus from Rousillon, Perpignan.
OOS ON MAMMALS FROM WESTERN JAVA. Bye
6. Ona Collection of Mammals from Western Java presented
to the National Museum by Mr. W. EH. Balston.
By Otupriznp Tuomas, F.R.S., F.Z.8., and R. C.
Wrovecuton, F.Z.8.*
[ Received February 27, 1909. |
Up toand during the earlier part of the last century no tropical
country in the world had more attention paid to its zoology than
the Island of Java, quite a considerable number of species being
described from there, firstly by the French, who received
specimens from Diard and Duvaucel, then by Dr. Thomas
Horsfield, who in 1824, under the auspices of the East India
Company, published a special work f on the subject, and finally
by the Dutch authors Schlegel and Miiller, who included many
Javan animals in their general work { on the Mammals of the
Kast Indian Archipelago (1839-44).
But from the latter date to the present time almost nothing
has been done, and but few specimens have been collected except
those that have gone to Leyden; and workers in the other
principal Museums have been often embarrassed by the absence
of good Javan specimens, representing the early described species,
for comparison with examples from other localities.
Of late, in the working out of Bornean and other Malay
material, the want of Javan collections has been severely felt, and
we are therefore proportionately indebted to Mr. W. E. Balston
for enabling the well-known collector Mr. G. C. Shortridge to
make a collecting-trip to Java after he had finished the work in
Western Australia of which Thomas had previously given an
account in our ‘Proceedings.’
Mr. Shortridge found the island extremely favourable for
collecting, and by the kind assistance of Mr. M. C. Kirkpatrick,
of the Hongkong and Shanghai Bank, and of the local Museum
authorities, he was enabled to visit many interesting localities and
to make one of the finest collections that we have ever received
from any one region.
In all, Mr. Shortridge obtained over 1500 specimens in the
island, the majority of which, however, were Bats, a group
in which Java is astonishingly rich. The species number 74, of
which we have found it necessary to describe six as new. In
addition, good series were obtained of a large number of species
which were either not in the Museum collections at all, or else
only represented by specimens from Dr. Horsfield’s collection,
which had been first to the East India Company’s Museum and
then transferred, mostly in 1879, to the British Museum. These,
while of extreme value as types or historical specimens, are
* Published by permission of the Trustees of the British Museum.
+ ‘ Zoological Researches in Java,’ 1824.
~ In Temminck’s ‘Verhandelingen over de Natuurlijke Geschiedenis der
Nederlandsche overzeesche bezittingen, 1839-44.
302 MESSRS. O. THOMAS AND R. C. WROUGHTON ON [ Mar. 16,
mostly too faded and deteriorated to be of much practical use
in making comparisons.
For the present magnificent addition to the National Collection
we have every reason to be most grateful to Mr. Balston, to
whom we were already so heavily indebted for his W. Australian
donations.
We have appended to the species Mr. Shortridge’s notes on
their habits, native names, and distribution.
1. HyLopatEs LEUcIscUS Schreb.
3. 492. @. 486, 487. Tji Wangie, Preanger.
‘* *Qoa-oo0a’ (Soendanese). ‘ Wau-wau’ (Malay).
‘* Local, apparently confined to the mountains of West Java;
where it occurs its presence is easily detected by its continuous
ery, which carries a long distance and of which its native name
is an imitation.”—G. C. 8.
2. PRESBYTIS PYRRHA Horsf.
Semnopithecus vel Presbytis maurus auctorum, nec Schreber.
Os Use. Os UO mii AOD mmm RE mn, 12m
gandaran, Dirk de Vries Bay.
6. 614. ©. 615, 616 imm., 617 imm., 749, 787, 788 imm.
Tjilatjap.
$. 475, 4938. 2. 474, 479, 480. Tji Wangie, Preanger.
5 SPA, iin. SPAY inary, Kalipoetjang, Tyi-Tandoei
River.
A comparison of this fine series, all black specimens, with the
co-types of Horsfield’s “ Semnopithecus pyrrhus” does not confirm
the conclusions as to the specific distinctness of the black and red
forms of the group published by Dr. Jentink*. The skull of
No. 22.4, one of the co-types of the red form, is exactly matched
by Mr. Shortridge’s No. 615, a black “mawrus,” in all the
characters mentioned by Dr. Jentink, namely the size of the teeth,
the degree of prognathism, the length of the palate, and the form
of the chin. And the series shows that there is a good deal of
variation in all of these characters. We are therefore compelled
to adhere to the usual conclusion (which is confirmed by
Mr. Shortridge’s note) that the red monkey named pyrrhus by
Horsfield is an abnormal red form of the usual black animal that
has gone by the name of maurus or maura.
But, so far as names are concerned, the specific term maura
Schreber, which has hitherto been used for this animal, should
give place to pyrrha. For the primary basis of Schreber’s
maura was the ‘* Middle-sized Black Monkey” of Edwards, whose
plate and description show that it was not this species at all,
but a West African Mangabey, possibly Cercocebus fuliginosus
EK. Geoff.
We do not think the evidence for the identification of Geoffroy’s
* Notes Leyd. Mus. xiv. p. 119 (1892).
1909. ] MAMMALS FROM WESTERN JAVA, 373
auratus (Ann. Mus. Hist. Nat. xix. p. 93, 1812) with this
monkey is sufficient to justify us in using the name. No exact
locality was given and the description might apply to any of the
forms with pale yellow young, which occasionally keep the
youthful pelage throughout life. Horsfield’s pyrrha, though
equally based on an abnormal specimen, is unquestionably this
species.
“¢ Tiutoong ’ (Soendanese and Javanese).
“Very plentiful. When first born the fur of the young is
bright rufous, but soon becomes black as it grows older.
“¢ Abnormal adult rufous individuals occasionally occur in which
the black coat has never been assumed.”—G, C. 8.
3. PRESBYTIS AYGULA L.
Semnopithecus vel Presbytis mitratus auctorum.
g. 482imm., 491. 9. 476. 481. Tji Wangie, Preanger.
Linneus* based his Simia aygula on a monkey described by
Osbeck + from Java, which can only have been the species to
which the name mitratus has been usually applied.
** “ Soereli’ (Soendanese).
“Very much more local than the last species. It seems, like
Hylobates, to be confined to the dense forests in the mountainous
parts of West Java. When very young the markings resemble
those of P. comata from Sumatra.”—G. C. 8.
4, MACACA FASCICULARIS Raff.
Cee Bataan
6.1777. &. 1721, 1780. Pangandaran, Dirk de Vries
Cu Olay" Oeics diene yoy
5. 490. 2. 470. Tyi Wangie, Preanger.
$. 1219. Tasikmalaja, Preanger.
3. 13879, 1428. Kalipoetjang, Tji-Tandoei River.
“«*Munyet’ (Malay). ‘ Keteq’ (Javanese) (Soendanese).
“Very abundant throughout Java.”—G. C.8.
5. Nycricesus JAVANIcuS E. Geoff.
6. 1371. Batavia.
“«¢ Mooka (Soendanese). Pookang (Javanese).
“ Nocturnal; not plentiful.”-—G. C. S.
6. Preropus vampyrus L.
Caen) Oe Zone bataiar
3S. 142. @. 148. Buitenzorg.
3. 795, 933. 2. 790, 794, 796, 797, 843, 934. Tjilatjap.
3. 1004, 1146, 1159, 1223. Tasikmalaja, Preanger.
* Syst. Nat. (10) i. p. 27, 1758, + Ostindisk Resa, p. 99, 1757.
wo
~
i
MESSRS. 0, THOMAS AND R. C, WROUGHTON ON [ Mar. 16,
. 1607, 1608, 1609, 1772; 1773, 1774, 1784, 1789, Pangan-
sae Dirk de Vries ee i
“¢ Karlong’ (of natives).
“Very plentiful except at high altitudes, especially where they
have formed colonies, Flight slow and flapping, occasionally
soaring like a bird. They seem to see fairly well by day, and
when ‘fyi ing will dip like a crow to avoid a bullet. In the
neighbourhood of their colonies there are generally a few flying
around all day, while those on the trees are continually altering
their positions and quarrelling among themselves, the peculiar
screaming that they make being audible for ‘a considerable
distance. When disturbed they soon take to flight, circling round
the trees like rooks, getting gradually up out of gunshot,
although eventually returning to the same place. Many
thousands generally collect in one colony, and the trees that they
have chosen (which are generally high ones) present a very
peculiar appearance, being almost entirely stripped of leaves,
while the bats hang in full view in thousands from all the upper
branches. ’—G. C. s.
. Rovserrus SHORTRIDGEI,
Thos. & W vought. Abstr. P. Z.8 SIRS ESAs
¢. 1905. Kalipoetjang, Tji-Tandoei R., 8. Java. 5 March,
1908. B.M. No. 9.1.5.67. — Z'ype.
Most closely allied to the continental 2. leschenaulti Desm. *,
with which it agrees in having the posterior lower molar elongate
instead of subcircular, as it is In amplexicaudatus Geoff. Size,
however, much greater, the skull especially being markedly
larger and heavier, and its crests more strongly’ developed.
Teeth throughout larger and broader, the anterior lower premolar
conspicuously larger than in the allied species.
Fur brown above, the hairs with lighter bases; hinder neck
nearly naked, paler; a paler patch over the base of the’ tail.
Below, the hairs are everywhere brown with lighter tips.
Dimensions of the type, the starred measurements taken in
the flesh :—
Forearm 94 mm.
*Head and body 145; .*tail 17; *ear 22; third finger,
metacarpal 60, first phalanx 39, second phalanx 51; lower
leg and hind foot (ce. u.) 66.
“Skull: aber length 42, zygomatic breadth 26-2, interorbital
breadth 8-9; supraor pital NRRL to tip of nasals 20:5 ; breadth
of ae 16:5; palate, breadth outside m* 13; palation to
incisive foramina 20:2; front of canine to back of m* 16:1,
length of m, 2.
Type as above.
This fine Rouset is the largest of the Oriental species, its
forearm exceeding those of ZR. ‘leschenaulti_ and. amplexicautatus
+ Cf. K. Andersen, Ann. Mag. N. H. (7) xix. pp. 501 et segq., 1907,
1909. ] MAMMALS FROM WESTERN JAVA. 375
by more than 6 mm. Its skull is large and heavy, and its
cranial ridges far more developed than in specimens of similar
age of the allied forms. In company with it there lives the
smallest eastern species, 2. minor, such a difference in size no
doubt tending to reduce competition, and therefore to enable
such closely allied forms to live side by side.
h. shortridgei must be exceedingly rare, as getting all the fine
series of bats now enumerated, including 34 of the previously
almost unknown ft. minor, My. Shortridge only obtained one
single specimen of it.
8. Rovuserrus mixor Dobs.
g. 1410, 1411, 1413, 1414, 1439, 1440, 1441, 1445, 1462,
1464, 1508, 1509.
2. 1409, 1412, 1442, 1443, 1444, 1463, 1465, 1506,. 1507,
1510, 1511, 1512.
Four males, six females in spirit.
Kalipoetjang, Tji-Tandoei R., 8. Java.
This species has not hitherto been represented in the Museum
collection, the only known example of it being the type, preserved
in the Calcutta Museum. The present fine series of topotypes
is therefore a most valuable accession to the National Collection.
“ «Tiddot’ (Javanese) (as distinguished from insectivorous
bats).
“The smaller fruit-bats are really much more plentiful than
Pteropus vampyrus, but on account of their smaller size and
late appearance in the evenings they are less noticeable. They
also seldom roost by day in conspicuous positions, but generally
collect in caves, hollow trees, or among dense foliage. When
disturbed by day they will dart off sharply in every direction
like insectivorous bats. The fruit-bats are a very serious pest
in Java, on account of their enormous numbers, and it is chiefly
on their account that so much of the fruit is picked green and
not allowed to ripen on the trees.”—G. C. 8.
9. CYNOPTERUS TITTHZCHEILUS Temm.
3.8, 12. Q.7,9,11. Low-lying country, Batavia. And
19 -m al,
3. 38,180. 9. 83. Buitenzorg. 850’.
g. 531,532. 2. 533. Svekaboemi. 2100’,
g. 684, 687, 689, 690,914. 2. 688, 691, 692, 693, 912, 913,
915,916. Tyjilatjap. Sea-level.
2. 1062. ‘Tasikmalaja, Preanger. 1145’.
The Buitenzorg specimens are absolute topotypes of this fine
- species, which had not previously been represented in the Museum
collection,
Young specimens of (1. titthecheilus are of a brownish-grey
colour, and in other respects have a general likeness to Rousettus,
for members of which genus they were at first mistaken.
376 MESSRS. 0. THOMAS AND R. C. WROUGHTON ON [ Mar. 16,
10. CYNOPTERUS HORSFIELDI Gray.
Ole 8299 MOO MOS LOS OSs 5 O 280 SoLSienos.
102, 104, 107, 109, 124. Buitenzorg. 850’.
3. 671, 778, 784, 818, 849, 861, 911. &. 777,835, 848, 944.
Tjilatjap. Sea-level. 1 ¢,6 2 inal.
2. 1650, 1652. Pangandaran, Dirk de Vries Bay.
6. 11386. @. 1042, 1061, 1137, 1288, 1289. Tasikmalaja,
Preanger. 1145.
We use provisionally Gray’s name for this Bat, as being unques-
tionably pertinent to it, but there are several earlier names, based
on specimens from Sumatra and elsewhere, which may hereafter
prove to be applicable to it. Why Mr. Glover Allen *, in writing
of Javan specimens, should use the name brachysoma Dobson,
when he knew of the 30 years’ earlier horsfieldi, we do not quite
understand.
The males of the Tjilatjap series are particularly brilliantly
coloured.
11, MaAcroGuossus minimus Geoff.
3d. 677, 678, 755, 773, 774, 775, 817, 827, 834, 844, 845, 880,
889, 891.
2. 670, 676, 700, 776, 785, 816, 832, 833, 890, 903. Tyilatjap.
Sea-level.
©. 1006, 1011. Tasikmalaja, Preanger. 1150’.
“<Tiodot-merah’ (Javanese). ‘ TjOd6t-pisang’ (Javanese).
“ Local, very abundant around Tyilatjap.
“Tongue highly extensile, but probably adapted merely for ex-
tracting the pulp from ripe fruit. None of the specimens
examined had traces of insects in their stomach. They may be
partly honey-eaters, Tail normally 2 to 3 millimetres in length,
but frequently entirely rudimentary externally.
‘¢ Fully adult specimens of both sexes have a peculiar crescent-
shaped gland on the lower part of the throat.”—G. C. 5S.
12. RHINOLOPHUS BORNEENSIS Peters.
@. 875. Tyilatjap.
3g. 1632. @. 1655, 1692. Pangandaran, Dirk de Vries Bay,
S. Java.
No member of this group of Lhinolophus had previously been
known from Jaya.
The posterior process of the nose-leaf is particularly short in
these specimens. of
“* Kampret’ (Malay). ‘Lalli’ (Soendanese). ‘Tyodot’
(Javanese) (generally only the smaller fruit-bats). ‘ Booerong-
tekoos’ (bird-mouse). ‘ Lawo’ (Javanese).
‘* Most of the larger imsectivorous bats are cave-dwellers, but
many of the smaller species hide by day in hollow bamboos,
* Bull. Mus. Harv. li. p. 25, 1908.
1909. ] MAMMALS FROM WESTERN JAVA. 3G
especially in those used in constructing the roofs of houses, and
occasionally in the rolled up young ieaves of bananas, hollow trees,
crevices among rocks, etc.’—G. C. 8.
13. RurnoLopHus AFFINIS Horsf,
36.514. ©. 510, 512, 513, 515, 517, 518. Kottamanah
Cave, Soekaboemi. 2100’.
oe 009) 2 256i (mi spirit); Do;
3. 696. Tyilatjap. Sea-level.
3. 1432, 1433, 1435. 9. 1434, 1436, 1437, 1461, 1470,
1471, 1472, 1473. Kalipoetjang, Tyji-Tandoei R., 8. Java.
6d and3 2 inspirit. Do.
The present fine collection adds considerably to our knowledge
of the Javan species of the genus Rhinolophus, for while in
Dr. K. Andersen’s important account of the genus* only six
species are recorded from the island (affinis, pusillus, acuminatus,
trifoliatus, luctus, and geminus), no less than three species are
now added to the list (viz. stheno, borneensis, and canuti)—thus
bringing the total up to nine.
14, RHINOLOPHUS ACUMINATUS Pet.
Gio He By Wea MO. WD, colin ters tos Oil yore) 1 Gin ally) deine
tenzorg. 850’.
Gy Ae PGR Oeil FS USO ISS Os aio: laine
(AS ont WAS Sa Sl SRS 4 seals, Tasikmalaja,
Preanger. 1150’.
ao OS in spirit Do:
With the exception of one more rufous individual, these
specimens are all dark smoky grey, while in afinis the usual
colour 1s more or less buffy. Some few individuals of the latter,
however, approximate to the colour of acwminatus.
It is noticeable that the two species were not obtained at any
one locality, as though they competed with each other too closely
to live in the same districts.
15. RurtnoLopHus stHENO K. Anders.
Oe) VLATG AOS: 2. 1429, 1475. Kalipoetjang, Tiji-
Tandoei R., S. Java.
Described by Dr. Andersen from the Malay Peninsula t, and
afterwards recorded by him from Sumatrat.
These specimens are all of a more or less buffy or fulvous
colour, while the skins of 2h. borneensis are uniformly dark grey.
Even on the dried skins Ah. stheno may be distinguished from
Rh. borneensis by the much longer posterior lancet of its nose-
leaf.
* Ann. Mag. N. H. (7) xvi. p, 656, 1905.
fe Ze SLO OSs aap oles
¢ Ann. Mus. Genoy. (8) iii. p. 24, 1907.
Proc. Zoou. Soc.—1909, No. XXV. 25
378 MESSRS. 0. THOMAS AND R. C. WROUGHTON ON [ Mar. 16,
16. RuarNoLopHus PusitLus Temm.
R. minor Horsf., nee Vespertilio ferrum-equinum minor Kerr
(1792), which is a Rhinolophus.
2. 504. Kottamanar Cave, Soekaboemi. 2100’.
2. 1236. Tasikmalaja, Preanger. 1145’.
@. 1477. Kalipoetjang, Tji-Tandoei R., 5. Java.
17. RHINOLOPHUS CANUTI.
Thos. & Wrought. Abstr. P. Z. 8. 1909, p. 18.
S$. 1481, 1474. 2. 1430, 1504. Kalipoetjang, Tji-Tandoei
River, 8. Java. 2 & 4 March, 1908.
Closely allied to 2A. creaght Thos., described in 1896 from
N. Borneo*, but distinguished by the following characters:—The
usual connecting-process between the sella and the posterior
lancet, obsolete (as a unique character) in ereaghi, is here repre-
sented by a narrow crest on the hinder side of the sella, less than
1 mm. in depth, and joining the posterior process low down ;
therefore very different from the usual high connecting-process,
between which and the obsolete one of creaghi it forms an inter-
mediate stage; the upper profile of the crest is evenly rounded
above and joins the top of the front face of the sella at a sharp
angle, without any intervening notch such as is usually formed
by the front surface of the sella projecting a little higher up-
wards than the anterior point of the connectiug-process. The tuft
of hair on the front face of the posterior process is very much
longer and more abundant, hiding the posterior end of the
connecting-process, and forming a most striking and prominent
feature of the bat; the hairs forming the tufts are shining
golden yellow, and cover the whole middle part of the posterior
leaf, some of them also going on the sides of the connecting-process
and on the back of the sella; there are also fine tufts of hair on
each side at the junction of the horseshoe with the outer bases of
the posterior process. The skull is essentially similar to that of
creaghi in the development and shape of the rostral projection,
which is, however, slightly higher, and has its front edge vertically
above the anterior part of p‘, while in creaghi that edge is over
the back of the same tooth, the canine-bearing part of the jaw in
front of the projection appearing therefore to be longer in creaght
than in canwée.
P' in the tooth-row; p, minute, external, p, and p, touching
each other internal to it; 1m one jaw out of eight it is absent.
Forearm 50 mm. Length of skull from occiput to front of
canine 22°5.
Head and body 65 mm.; tail 22; ear 24.
Hab. as above.
* Ann. Mag. N. H. (6) xviii. p. 244. In the eighth line of this description, for
the word “ sella” read “ posterior process.” The second lower premolar (ps), said to
be “entirely absent,” proves, on the skull being cleaned, to be present on one side,
though excessively minute.
1909. | MAMMALS FROM WESTERN JAVA. 379
Type. Adult male. B.M. No. 9.1.5.183. Original number 1431.
Collected 2 March, 1908.
We have named this striking species of Rhinolophus, which con-
nects the aberrant /. creaghi with the more normal species of the
genus, in honour of Dr. Knud Andersen, in recognition of the
exhaustive work he has done on this complicated and difficult
group.
18. HipposipEros piaADEMA Geoff.
3. 1126, 1147, 1302, 1303. 9. 1296, 1301. ‘Tasikmalaja,
Preanger. 1150’,
3. 1383, 1403, 1406, 1407, 1422, 1423, 1494,1515. 9. 1405,
1421, 1424, Kalipoetjang, Tji-Tandoei R., 8. Java.
3. 1603. Pangandaran, Dirk de Vries Bay, 8. Java.
representative of HZ, wobilis Horsfield.
19. Hrpposiperos prcotor Temm.
2. 499, 500, 505. Kottamanah Cave, Soekaboemi. 2100’.
9 inal. Tyilatjap.
20. Hiprosrperos LArvAtus Horsf.
3. 1385, 1586, 1391, 1446. 2. 1384, 1387, 1388, 1389, 1390,
1392, 1447, 1448. Kalipoetjang, Tji-Tandoei R., 8. Java.
3. 1604, 1647, 1653, 1674, 1675, 1679. 9. 1605, 1648,
1649, 1677, 1678, 1680, and 10 specimens (4 3, 6 2) in spirit.
Pangandaran, Dirk de Vries Bay, 8. Java.
The forearms of these specimens run from 55 to 59 mm. in
length. Dobson gives 2°45 in. (=62°5 mm.).
Of the 24 individuals in skin, four (all females) are strongly
suffused with yellowish.
21. CaLops FrirHit Bly.
3. 1024, 1025, 1026. Tasikmalaja, Preanger. 1145’.
Topotypes of C. bernsteini Peters.
These examples of this exceedingly rare Bat are particularly
welcome, as representing Peters’s C. bernsteini, which is not im-
probably distinguishable from Blyth’s C. frithii. But pending
further investigations we use the name adopted by Dobson.
The recently described (C. robinsoni Bonh., from Selangore, is
much smaller than either.
92, MEGADERMA SPASMA TRIFOLIUM Geoff.
g. 74. Buitenzorg. 850’.
3. 853, 856, 863, 866, 870, 874, 878, 894. 9. 695, 739, 854,
855, 857, 858, 865, 867, 868, 869, 871, 872, 873, 879, 892, 893.
Tjilatjap.
g. 1155. Tasikmalaja, Preanger. 1150’.
25*
380 MESSRS, 0. THOMAS AND R. C. WROUGHTON ON [Mar. 16,
23. PETALIA JAVANICA Geoff.
@. 81,90, 97. Buitenzorg. 850’.
3. 643, 649, 650, 651, 652, 655, 656, 657, 658, 659, 662.
©. 642, 644, 645, 647, 653, 654, 661, 663. Cave on sea-coast,
Tjilatjap. And1 g and 11 9 inal.
6. 1029, 1030. @. 1027, 1028. Tasikmalaja, Preanger.
Although the general colour of these specimens varies con-
siderably, from grey to fulvous, yet the latter colour is no more
dominant or intense in the males than in the females, as might
perhaps have been expected.
Dobson’s record of ‘‘ Timor” for an example of this species is in
all probability erroneous, as the specimen was purchased from a
dealer, and the entry of its locality in the register 1s somewhat
ambiguous.
. TYLONYCTERIS PACHYPUS Temm.
24
g. 41,42. ©. 94,132. _Buitenzorg. 20 ¢ and 42 9 inal.
6. 520,527. ©. 521, 526,528. Soekaboemi.
3. 993, 995, 999, 1093. @. 978, 980, 982, 996, 997, 998,
50, 1078, 1140. Tasikmalaja.
95. PIPIstRELLUS TRALATITIUS Horsf.
©. 5,224. Batavia.
6. 126,185. @. 61, 181, 182, 183,184. Buitenzorg.
2. 530 and 18 specimens in al. Soekaboemi.
3. 598, 600, 602, 605. &. 597, 599, 601, 606, 608, 609, 610,
- 611, 626, 627, 697, 698. 10 9 inal. Tyilatjap.
3. 1000, 1001, 1229. @. 1035, 1055, 1056, 1200. Tasik-
malaja.
This is the Bat for which Dobson uses the name abranus
Temm., although Horsfield’s name is earlier and one of the
specimens the former enumerates (w’) he states to be labelled
“ Lowo-manir,” the native Javanese name recorded in Horsfield’s
original description. Specimen A of Horsfield’s ‘ Catalogue of the
Indian Museum’ (p. 39), now B.M. No. 79.11.21.124, may be con-
sidered as the type of tralatitius.
As elsewhere in the East, this Bat seems to be excessively
common in Java.
26. PIPISTRELLUS IMBRICATUS Horsf.
@. 756, 757, 758, 760. Tyilatjap.
27. ScoroPHILUS TEMMINCKII Horsf.
Go oo, Gr, GO, WU. iis, WO. Man May, Wee, Wad Idi 1s.
©@. 68, 113, 114, 115, 116, 119, 129, 159. Buitenzorg. 850’.
3. 225. Batavia. :
g. 524. ©. 519, 525. Soekaboemi.
So Gli, SOG, Gls, SA), sO, SS, se, ose, GAO, O@. Ob,
807, 808, 809, 827, 828, 836, 927, Tyilatjap.
1909.] MAMMALS FROM WESTERN JAVA. 381
a 902, LOOT LIGl, 1207 Oy 1092) 1204 sI2067 9 Dasik-
malaja, Preanger. 1150’.
We use the name temminckii for these bats, as being un-
doubtedly pertinent to them, pending such time as the type of
kuhli can be definitely assigned to one or other form of Scoto-
philus.
28. MuRINA BALSTONI Thos.
Ann. Mag. N. H. (8) ii. p. 370. 1908.
@. 1160. Tasikmalaja, Preanger, 1145. B.M. No. 9.1.5.354.
Type.
This, the first new species distinguished of the fine Javan
collection made by Mr. Shortridge, was named by Thomas after
Mr. W. EH. Balston, to whom the National Museum owes this
magnificent addition to its collections.
The distinctive characters of J/. balstoni are enumerated in the
paper above referred to.
29, HARPIOCEPHALUS HARPIA Temm.
3. 901, 902. ©. 899, 900. Thilatjap. 10 Dec., 1907.
2. 1287. ‘Tasikmalaja, Preanger. 1145’. 28 Jan., 1908.
Compared with these specimens the Himalayan form is obviously
different by its darker and more chocolate colour. It should bear
the name of H. lasyurws Hodgs.*, of which the Museum specimen
No. 79.11.21.119 is the type. Horsfield’s Lasiwrus pearsoni, also
from Darjiling, would be a synonym of it.
30. Myoris sp. (near MURICOLA).
Sioa 00, at, W405 O29 30. oo and 19) OF my, al,
Buitenzorg.
3. 803, 885, 887. ©. 802, 804, 886, 925. Tjilatjap.
3. 977,1162,1191,1193. ©. 1139. Tasikmalaja, Preanger.
31. Myotis (Leuconor) HAsseLti Temm.
g. 1017, 1054, 1069, 1070, 1071, 1072, 1073, 1090, 1100,
HOI WO2 eNOS eNO aT LOGs Mbt tel) Tiiee std. Toe)
1194. @. 1074, 1105, 1107, 1108, 1109, 1110, 1132. Tasik-
malaja, Preanger.
32. Myoris (LEucoNogE) apversus Horsf.
2. 1621, 1622, 1623, 1637, 1638, 1639, 1640, 1643, 1651,
1656, 1657, 1658. @. 1614, 1615, 1616, 1617, 1618, 1619,
1624, 1633, 1641, 1644, 1645. Pangandaran, Dirk de Vries Bay.
33. KeERivouLa Prcta Pall.
@. 819, 820. Tyilatjap. Sea-level. 27 Nov., 1907.
““« Lawo-Kambang’ (Javanese).”—G. C. S.
* J. A.S. B. xvi. p. 896, 1847.
382 MESSRS. 0. THOMAS AND R. C. WROUGHTON oN [ Mar. 16,
34. KERIVOULA HARDWICKEI Horsf.
3g. 14,16. Batavia (in spirit).
6 A065, Vs se TOP OCS. M064 LOGGAOG 77 10687 1125,
1148. Tasikmalaja, Preanger. 1145’.
Dobson’s Catalogue measurement of 1-4 in. for the forearm of
K. hardwickei is more than any of these specimens show, as they
run from 31-33 mm., therefore only rarely reaching 1°3 in., which
was the size of his A. fusca, later synonymized by him with
K. hardwicker.
35. MINIOPTERUS BLEPOTIS Temm.
go. 1396, 1398, 1399, 1500, 1518, 1519, 1533, 1534, 1536,
1537. 9. 1393, 1394, 1395, 1397, 1400, 1401, 1402, 1516,
1517, 1535. 9 Qand1 ginal. Kalipoetjang, Tyi-Tandoei R.
g. 1152. 2.1151. Soekaboemi, Preanger.
Topotypes. Temminck gives length of forearm as 45:5 to 47°5;
the present series vary from 44:5 to 46, but those of the two
specimens from Soekaboemi measure 50 mm.
36. MINIOPTERUS MEDIUS, Sp. n.
gd. 1498. ©. 1497, 1499, 1501, 1502. And 1 ¢,2 9 imal.
Kalipoetjang, Tji-Tandoei R.
A Miniopterus intermediate in size between JV. blepotis and
M, tibialis.
Not differing materially in outward facies from JV. blepotis,except
in size. Black and red phases of colouring of the coat present as
is usual in the genus.
Dimensions of the type, the starred measurements taken in the
flesh :—
Forearm 42 mm.
Head and body *55; tail *47; hind foot *9; ear *12; terminal
phalanx of middle finger 35.
Skull: greatest length 15; basi-sinual length 11:5; brain-
case breadth 75; greatest width across canines 6°5; front of p*
to back of m?® 4°4.
Hab. Kalipoetjang, Tji-Tandoei R., Java.
Type. Adult female. B.M. No. 9.1.5.464. Original number
1499. Collected 4 March, 1908.
37. MINIOPTERUS TIBIALIS Tomes.
3S. 1451, 1453, 1454, 1456, 1457, 1458, 1483, 1486, 1492.
@. 1449, 1450, 1452, 1455, 1459, 1460, 1481, 1482, 1484, 1485,
1487, 1488, 1489, 1490, 1491. And 5 Q in al.
38. EMBALLONURA MONTICOLA Temm.
@. 1514. Kalipoetjang, Tji-Tandoei R., 8. Java.
3. 1602, 1628, 1629, 1634, 1636, 1664, 1672. °. 1600, 1601,
1612, 1630, 1635, 1659, 1660, 1661, 1662, 1663, 1665, 1666, 1668,
1909. | MAMMALS FROM WESTERN JAVA. 383
1669, 1670, 1671, 1673. Pangandaran, Dirk de Vries Bay, 8.
Java.
Most of these specimens are rather larger than is indicated in
Temminck’s description, which gives a forearm length of “ 1 duim
7 lijnen” (=43 mm)*. Their forearms range from 43°5 to
48 mm.
39. TAPHOZOUS LONGIMANUS Hardw.
¢o.6. Batavia. Sea-level.
6. 672. Tijilatjap. Sea-level.
These specimens are paler than usual, but this may be due to
bleaching, or to a variation of the same nature as that found in
Blyth’s “ fulvidus.”
40, TapHozovus THEOBALDI Dobs.
6. 1415, 1417, 1418, 1419, 1420, 1469, 1479, 1480, 1496.
@. 1416, 1426, 1478, 1495. Kalipoetjang, Tji-Tandoei R., 8.
Java.
3 56,5 2 in spirit. Do.
This rare Bat was hitherto only known from the typical speci-
mens in the Calcutta Museum, of which one has been given to
the British Museum, and has enabled us to compare Mr. Short-
ridge’s specimens with it. We can find no essential difference
between the Javan and Tenasserim forms, although when fresh
skins of the latter are available some difference in colour may
prove to exist.
As represented by the Javan specimens, 7. theobaldi has a
blackish throat, recalling the black beard of 7’. melanopogon, but
the black is not so deep nor so sharply defined laterally as in that
animal.
41. TAPpHozous sACCOLAIMUs Temm.
2. 1363. Tasikmalaja, Preanger.
42, CHG@REPHON PLICATUS Buch.-Ham.
3. 58, 59, 62,112. 9.57. Buitenzorg. 850’.
3. 535,537. ©. 508, 534, 536. Soekaboemi.
3. 946, 947, 948, 949, 950, 952, 954, 955, 956, 957, 971.
@. 951, 953. Tyilatjap.
@. 1290. Tasikmalaja, Preanger. 1150’.
And1¢,5 @ inal. Batavia.
Represent Horsfield’s Vyctinomus tenuis and dilatatus.
The Tjilatjap specimens Nos. 946-957 are all quite young.
“Very active on the ground, and even when young and unable
to fly is able to run about like a mouse, especially in rough situa-
tions.” —G. C. 8.
* Not 40 as stated by Miller, who has taken Temminck’s measurement as though
the inches were English ones instead of French. As a consequence, there is no
difference at all in size between H. monticola and EF. peninsularis Miller.
384 MESSRS. 0. THOMAS AND R. C. WROUGHTON ON [ Mar. 16,
43. GALEOPTERUS VARIEGATUS Geoff.
3. 213. Batavia.
6. 120, 2 2 oiler io, eneG. Ox, TUGIO:
1716, 1717, 1724, 1730, 1778, 1779 imm. Pangandaran, Dirk de
Vries Bay.
“«* Tando’ (Soendanese). ‘ Walang-kopo’ (Javanese).
“Nocturnal. This animal possesses a very strong and peculiar
smell, which originates from an open gland at the root of the tail.
This gland is in the form of a shallow pouch—more developed
in the males—and, during life, a vivid orange in colour. The fur
of the normal-coloured variety of the species has a distinct suf-
fusion of green during life that at once disappears when the skin
is dry. This peculiarity is also noticeable in 7upaia.”—G, C. 8.
Ad, TUPAIA JAVANICA Horsf.
3g. 148, 174, 175, 189, 247, 248. 9. 51, 52, 156, 246.
Buitenzorg.
Go (50, 2, fds Ainlkyaeyo.
3. 485. 9.473. Tyi Wangie, Preanger.
3. 1009, 1010, 1023, 1046, 1047, 1052, 1094, 1130, 1167,
1210, 1211, 1213, 1239, 1240, 1248, 1254,1292. ©. 1018, 1019,
1020, 1021, 1037, 1053, 1075, 1157, 1158, 1166, 1168, 1169,
1222, 1227, 1241, 1247, 1253. ‘Tasikmalaja, Preanger.
“ « Kekkas’ (Soendanese). ‘ Emess.’
‘“¢ Diurnal—bearing an extraordinary resemblance to the smaller
squirrels both in habits and movements; frugivorous and insec-
tivorous.”—G. C. 8.
45. CROCIDURA sp.
@. 45. Buitenzorg.
46. PACHYURA MURINA.
©. 13. Batavia.
3. 47, 63, 75, 79, 147, 163, 179, 188. &. 28, 46,56, 64, 144,
187, 190, 203. Buitenzorg.
3. 881. Tyilatjap.
“«¢ Tekoos-tjeroot’ or ‘Chuleroot’ (Jav., Soend., Mal.).
“ Probably originally introduced.
‘“‘Both species of Shrew seem to be most plentiful in thickly
populated localities, especially around seaport towns.”—G. C. S.
A7, Feuis parpus L.
1307 (black variety). Tasikmalaja, Preanger.
1156, ¢ 1294. Tasikmalaja, Preanger.
1260. Pangandaran, Dirk de Vries Bay.
‘“¢¢Matjan-tootool’? (Soend., Jav.). ‘ Matjan-kambang’ (black
var.).
“Comparatively plentiful throughout Java, particularly in
mountainous and thinly populated districts.”—G. C. 8.
1909.) MAMMALS FROM WESTERN JAVA. 385
48. FELIS JAVANENSIS Desm.
239, 1328, 1329imm., 1330. Batavia.
S$. 53 imm., 110 1mm., 171, 855 imm. Buitenzorg.
3. 1685 imm., 1686 imm. Pangandaran, Dirk de Vries Bay.
6.791. Thilatjap.
6. 1165, 1233 imm.,, 1245 imm., 1364 imm. @. 1257 imm.,
1367 imm., 1368 imm. Tasikmalaja, Preanger.
“«« Meong-chongkok ’ (Soendanese). ‘ Blatjan’ (Javanese).
“Very plentiful. Appears to occasionally cross with the
domestic cat.
“The domestic cats in Java almost invariably have distorted
tails, which are either stunted or have the appearance of being
disjointed, often in several places, the end being sometimes almost
at right angles with the base; very few, except recently imported
individuals, possess perfect tails.”—G. C. 8.
49, VIVERRICULA RASSE Horsf.
18, 230, 231, 233 imm., 234, 236. ©. 3/7, 227. Batavia.
164,167imm. @. 39,169. Buitenzorg.
3. 741 imm., 742 imm., 847imm., 970 imm. = Tyilatjap.
6. 1127, 1164, 1242, 1284 imm., 1285imm., 1286 imm., 1293,
1298. ©. 1058, 1209, 1224, 1244, 1297 imm., 1299 imm.
Tasikmalaja, Preanger.
‘¢ Rassie’ (Malay). ‘ Deddess’ (Soendanese).
“Very plentiful. It is curious that so many of the native
mammals in Java, especially the small carnivores, abound in the
densely populated districts. Viverricula, Paradoxurus, Mungos,
Felis javanensis, etc., seem to have entirely adapted themselves to
an artificial, almost semi-domestic life, existing in abnormal
numbers around towns and native villages, where they probably
live almost exclusively on the swarms of house-rats and poultry
that are always in such numbers in the vicinity of houses.
Viverricula is often kept alive, or hunted for by the natives, who
collect its civet either as a perfume or for flavouring their tobacco.
“Tt is astonishing how many even of the carnivorous mammals
of Java are partly frugivorous. In fact, there are few, if any,
mammals of any kind there that will not feed on bananas.’—
GVers:
50. PARADOXURUS JAVANICUS Horsf.
3. 43, 162,194. 9. 44, 55, 70, 131, 202. Buitenzorg.
1331, 1332, 1333. Batavia.
3. 1525. Kalipoetzang, Tji-Tandoei River.
3. 1783 (imm.). @. 1706. Pangandaran, Dirk de Vries Bay.
2. 935. Tyilatjap.
6.) 1133 imme, 34 imme: M63. (258hmm 2) LO9Sammr,,
1150 imm., 1153 imm., 1154 imm., 1214 imm., 1251 imm.,
1259 imm., 1260 imm., 1369 imm., 1370 imm. ‘Tasikmalaja,
Preanger.
386 MESSRS. 0. THOMAS AND R. C. WROUGHTON ON [ Mar. 16,
““* Looark’ (Jav.). ‘ Tjareuh’ (Soend.).
“ Very plentiful, largely frugivorous. Known in Java as the
Coffee-Cat, on account of its habit, when in the neighbourhood of
coftee- plantations, of feeding lar gely on the hen vias. The un-
digested berries, which are e afterwar ds dropped, are found in heaps
and carefully collected by the natives, and as the animals pick
out the ripest and best fruit, Looark-coffee is considered the finest.
‘“¢ Around houses it is often considered a useful factor in keeping
down the enormous numbers of house-rats that overrun so many
parts of Java; it is, however, frequently very destructive to
poultry. Often hides by day in the roofs of houses or out-
buildings. Although its general scent resembles that of Viverri-
cula, its gland-pouch is more or less rudimentary and does not
produce civet.”—G. C.8.
51. Cuon sAvanicus Desm.
6. 150imm. Buitenzorg Museum.
“¢ Adjac.’ ‘Garong’ (Soendanese). ‘ Andjing-o0tan ’ (Malay).
‘‘ Confined to the more inaccessible and mountainous parts of
Java. Said to be fairly numerous among the mountains of Hast
Java.
“| think that the native Campong-dog has most probably at
least partly originated from this species, and that C. yavanicus
will, like the ines of Australia, occasionally cross with domestic
varieties.” —G. ©. 8.
52. ARCTICTIS BINTURONG Raff.
1308. Near Tasikmalaja.
“¢ Matjan-thongkok.’ ‘Saro-garlong.’
“ Apparently rare in Java. Said to frequent the banks of
rivers.” —G. C.S8.
53. Muneos sAvanicus HE. Geoff.
232, 1320, 1321, 1822, 1323, 1324, 1325, 1326. ¢. 228.
Batavia.
3. 7limm., 191 imm.,192imm. Buitenzorg.
3. 896. 9%. 846imm. Tjilatjap.
3. 1304. o 1235, 1250, 1283, 1291. Tasikmalaja, Preanger.
“¢Garangan’ (Jav.). ‘Ganggarangan’ (Soendanese).
‘* Fossor ial; principally nocturnal; very plentiful.”—G.C.S.
54. HELICTIS ORIENTALIS Horsf.
153, 154. Buitenzorg Museum.
3. 205. Buitenzorg.
3. 1234imm. @. 1225, 1243. Tasikmalaja, Preanger.
“¢¢ Beool ’ (Soendanese).
‘“* Nocturnal and fossorial. This species has no offensive smell.
Probably mimics J/ydaus in its coloration, and like that animal
it frequents mountainous localities.”— G.C.S.
1909. ] MAMMALS FROM WESTERN JAVA. 387
55. MyDAUS JAVANENSIS Desm.
483. 92.478. ‘Tji Wangie, Preanger.
o. 149) immi 152) 168,09 178) 197 @., 145, 195, 200.
Buitenzorg.
The specific name meliceps, given by F. Cuvier, Mamm.
pl. 129, April 1821, is antedated by javanensis Desmarest,
Mamm. p. 187, 1820; and as both were based on a specimen
collected by Leschenault de Latour, there can be no question as
to their identity.
“¢ Seooong’ (Soendanese).
‘‘ Nocturnal and fossorial. Rather sluggish in its movements.
Its offensive smell, especially when disturbed ox irritated, can
hardly be exaggerated. Plentifulin the mountainous districts of
West Java.”—G.C.S.
56. Luvrra CINEREA III.
3. 1684. ©. 1682. Pangandaran, Dirk de Vries Bay.
3. 34 imm., 77 imm., 170 imm. @. 33 imm., 78 imm.
172imm. Buitenzorg.
1314imm. Batavia Museum.
g. 25imm. Batavia.
“« Saro’ (Javanese; Soendanese). ‘ Linsang’ (Jav.).
“Nocturnal; plentiful in swampy places and along the sea:
shore.”—G.C.58.
57. PETAURISTA NITIDA Desm.
157,158. Buitenzorg Museum.
A77. Tji Wangie, Preanger.
“¢ Belook’ (Soend.). ‘ Kooboong.’
“Nocturnal; apparently local. Said to be fairly plentiful in
some districts.” —G. C.8.
58. Scruroprerus (Hytoperes) sacirra L
S. lepidus Horsf., auct.
3. 897. 9. 898. Maos, near Tyilatjap.
3. 1626, 1627, 1790. @. 1625, 1791. Pangandaran, Dirk
de Vries Bay.
g. 1427. Kalipoetjang, Tji-Tandoei R.
We think we may venture now definitely to determine
Linneus’s Sciurus sagitta, which has been so long a puzzle to
workers on this group, its name having been assigned to quite a
number of different forms. Its locality is stated to be Java, and in
that island there are only two Flying Squirrels of which the phrase
“ statura Sciuri vulgaris” could be used, namely, S. lepidus Horsf.
and S. genibarbis Horsf., and from the latter of these it is
separable by “seéa una in mala,” sufficiently applicable to the
present animal. One statement only is inconsistent with this
conclusion, namely “ cwtis extensa a capitis ad carpum,” as though
there were an antebrachia! membrane as in the genus Petawrista.
388 MESSRS. 0. THOMAS AND R. C. WROUGHTON ON [ Mar. 16,
But as the specimen was clearly adult (‘ scrotwm magnum”) it
could not have been a young Petaurista nitida, and we can only
suppose that there was some error of observation in the matter of
the antebrachial membrane.
S. sagitta is a Hylopetes nearly allied to S, awrantiacus, spadiceus,
and everett.
‘““* Mon-mon’ (Soendanese). ‘Choobok’ (Javanese). ‘ Entjang-
entjang.’
“ Sciuropterus and Petaurista are nocturnal, while all the other
Javanese Squirrels seem to be diurnal.
‘<8. sagitta chiefly occurs among cocoanut-plantations, and either
makes its nest in an empty cocoanut or among leaves at the top of
a palm.”—-G. C.S8.
59. RATUFA BICOLOR Sparrm.
155 (juv.). Buitenzorg Museum.
g. 1381, 1382, 1425, 1520, 1522, 1526, 1529, 1530. ©. 1378,
1513, 1527, 1528, 1531. Kalipoetjang, Tji-Tandoei River.
3. 1698, 1710, LAE IS ON 1683, 1722, 1725, 1726.
Pangandaran, Dirk de Vries Bay.
3. 1295. ©. 1057. Tasikmalaja, Preanger.
“¢ Badjing-djalarang’ (Soend.) (Jav.).
“Tn uncultivated forest-districts ; local, often very plentiful
where it occurs.” —G. C.S.
60. Sciurus notatus Bodd.
. 1,3 Gmm.), 15 Gmm.). .9. 2,19. Batavia.
. 31, 49, 54, 245. Buitenzorg.
. 1468. 2. 1532. Kalipoetjang, Tji-Tandoei River.
. 1613. Pangandaran, Dirk de Vries Bay.
603, 612 624, 625, 629, 630, 631, 680, 801 (imm.).
©. 607, 619, 623, 632, 633, 681. Tjilatjap.
Examination of this series proves that Bonhote’s S. andrewst *
from Tyjigombong cannot be separated from S. notatus.
««« Badjing-klapa ’ of natives.
“ Very abundant among cocoanut-plantations in low-lying
country. In the mountainous and uncultivated localities its place
is taken by Sciwrus nigrovittatus. Although feeding on most
fruits, it seems to be especially partial to the cocoanut-palm, to
which it is very destructive. When feeding on a cocoanut it
bores a circular hole in one side of the shell. Natives frequently
fasten a piece of sheet tin around the trunks of the palms about
halfway up, as the tin offering no foothold prevents the squirrels
from ascending to the fruit.”—G. C.8.
Oe S230
61. ScruRUS NIGRO-viTTATUS Horsf.
3. 36 (imm.), 48, 186 (amm.), 244, 469. Buitenzorg.
2. 1380, 1503. Kalipoetjang, Tji-Tandoei R.
* Ann. Mag. N. H. (7) vii. p. 456, 1901.
1909. } MAMMALS FROM WESTERN JAVA. 389
¢. 618, 747, 751, 752, 753, 754, 780,781. 2. 635, 748, 782,
789. Tyilatjap.
Q. 471, 472, 488, 489 (imm.). Tji Wangie, Preanger.
3. 743,745. 9. 744, 746. Noesa Kambangan Island.
“«¢ Badjing-ootan.’
“Very plentiful in all the forest-districts. Unlike S. notatus
it avoids cultivation. Partly insectivorous.”—G. C.5.
62. LARISCUS * INSIGNIS JAVANUS.
Laria insignis javana, Thos. & Wr. Abstr. P. Z. Sh SOY joe WD:
¢. 65. Buitenzorg.
3g. 484. Tji Wangie, Preanger.
A Lariscus of the same general pattern as LZ. insignis, but rather
larger and stouter, and darker in coloration than any of the races
of that species.
Size rather larger than L. insignis of Sumatra. Fur about
15 mm. in length on the back, 30-35 mm. on the tail. General
colour-pattern as in the various races of L. insignis, but differing
from all of them in the darkness of the interspaces between the
dorsal stripes. Individual hairs of tail black, with a buff band
(5 mm. broad) at 5 mm. from the base, and with a white tip
(5 mm.); the outer black band (15 mm.) is much broader than in
other forms of Z. insignis, in which it measures at most 10 mm.
Skull with a long pointed muzzle, palatilar length 24 mm. as
compared with 19-22 mm. in Z. insignis and its Malayan
forms. ‘Teeth larger and stouter. Profile of skull much flatter.
Dimensions of the type :-—
Head and body 220 mm.; tail (broken) ; hind foot 47; ear 18.
Skull: greatest length 53; basilar length 41 ; zygomatic
breadth 28; interorbital breadth 14; palatilar length 24;
diastema 14; upper tooth-series, exclusive of p* 9°3.
Hab. Java (type from Buitenzorg). Alt. 855’.
Type. Adult male. B.M. No. 9.1.5.718. Original number 65.
Collected 2 August, 1907.
The second specimen, an old male, from Tji Wangie, Preanger
(alt. 4000’), agrees with the type in all essential characters.
‘he dark dorsal area, the broad black longitudinal stripe of the
tail, the long narrow muzzle, and large stout teeth serve easily
to distinguish this subspecies from typical LZ. insignis and its
Malayan races, peninsule and jalorensis.
“ ¢ Bo-oot’ (Soendanese).
‘‘ Mountainous districts.” —G. C. 8.
63. Nannoscrurus weLanotis Mill, & Schl.
3. 208. 2. 209. Buitenzorg.
* Nom. nov. Laria Gray, 1867, nee Scopoli, Entomol. Carniol. p. 21, 1763.
Type L. insignis.
390 MESSRS. 0, THOMAS AND R. C. WROUGHTON ON [ Mar. 16,
64, CHrROPODOMYS ANNA.
Thos. & Wrought. Abstr. P. Z.8. 1909, p. 19.
O01 03s Om oun eae re Soekaboemi.
. 674, 860, 932. 9. 799, 822. Tyilatjap.
1212, 1237, 1256, aah @. 1128, 1129, 1246, 1306.
Da cikemaleyas Preanger.
LOT 93) avar
A Chiropodomys outwardly resembling C. gliroides, but with a
smaller skull and teeth.
Size intermediate between the Bornean C. major and C.
pusillus. Fur rather short (6-7 mm. on the back), but close and
fairly soft.
General colour above between “isabella” and “fawn,” the
individual hairs being ‘ slate-colour” with “russet” tips; under
surface pure white, the hairs white to their bases. Hands and
feet whitish, the latter with dark central line above. Tail nearly
half as long again as the head and body.
Dimensions of the type (the figures in brackets are those of a
specimen of gliroides of similar age from Tenasserim) :—
Head and body 87 mm.; tail 112; hind foot 18; ear 16.
Skull: greatest length 24 [26]; basilar length 8°5 [20-5];
zygomatic “preadth 14 [15]; brain-case breadth 11:6 [13];
diastema 6°3 [7:3]; palatal foramina 3 [4:3]; upper molar
series 3°8 [4].
Hab. Tjilatjap, Java (sea-level).
Type. Adult female. B.M. no. 9.1.5.757. Original number
822. Collected 27 Nov., 1907.
Highteen specimens ex xamined, of both sexes and all ages.
In many respects this species is intermediate between the
mainland C. gliroides Bly. (syn. peguensis Bly., penicillatus Pet.)
and the Sumatran C. niadis Miller.
es Tékoos-klapa, Tekoos-sarwa, Ny ing-N ying.
“ Arboreal; similar in habits to alg Jlomaamse. making small
grass nests among bamboos or the heads of palm-trees.
“Very plentiful in many localities; seems to feed to a large
extent among rice-fields.”—G. C. 8.
Bio, 05
35. Mus rarrus (neglectus group).
Ge Ite ie ae sBatavias
g. 40, 50, 84,199. 2. 32, 122,133. Buitenzorg.
3. 628, 638, 641, 682, 738, 9386. 9. 639, 640, 664, 665, 821,
883, 884, 943. Tyilatjap.
d. 1115, 1201, 1202, 1208, 1252, 1262, 1264, 1266, 1267, 1268
1269, 1270, 1271, 1272, 1274, 1275, 1276, 1277, 1278, 1372, 1374,
1375. 2. 1022, 1040, 1041, 1220, 1249, 1261, 1263, 1265, 1273.
Tasikmalaja, Preanger.
3. 1538. Kalipoetjang, Tji-Tandoei River.
“<Tekoos’ of natives.
“ Swarming throughout Java, particularly in thickly populated
localities.” —G. C. 8.
1909.] MAMMALS FROM WESTERN JAVA. 391
66. Mus sp. (concolor group).
g. A495, 497. 2. 494, 496, 498,
67. Mus muscutus Linn.
2. 30, 73,141. Buitenzorg.
3. 646, 864. Tijilatjap.
3. 583. 9. 584. Soekaboemi.
3. 1279, 1280, 1281, 1282. Tasikmalaja, Preanger.
68. Banpicora serrrera Horsf,
3.1051. Tasikmalaja, Preanger.
3.1611. Pangandaran, Dirk de Vries Bay.
The first specimens of the Javan Bandicoot received since the
arrival of the original co-type, a faded and deteriorated skin
transferred from the Indian Museum in 1879.
Hermann’s Mus javanus might have been supposed to have
been this species were it not-for his distinct statement that its
feet were white, while in &. setifera they are dark brown. There
is, I think, little doubt that JL javanus was based on an example
of Mus norvegicus.
“<ekoos-djantang.’ ‘Tekoos-ia.’ ‘ Tékoos-besar,’
“Said to prefer swampy localities and to live in holes in the
ground, often among rice-fields.”——G,. C. 8.
7
69. Hysrrix savAnica F, Cuvy.
237, 1310, 1311, 1312, 1313. Batavia.
2.176. Buitenzorg.
2. 669. Tyilatjap.
9. 1631. Pangandaran, Dirk de Vries Bay.
“<< Tandark’ of natives. :
“Nocturnal and fossoria]l ; plentiful in many localities, said to
do considerable damage among rubber-plantations.”—G. C. 8.
70. Lerus nicricoLuis F, Cuv.
240, 241, 242, 1315, 1316, 1317, 1318. 9. 243. Batavia.
gd. 146,207. 2. 177imm., 204 imm., 206 imm. Buitenzorg,
“¢Klintji’ (Malay).
“Very local; apparently confined to the north-west of Java.
Plentiful among sugar-cane plantations around Batavia and
Buitenzorg. Probably originally introduced ; said to be plentiful
in Sumatyra.”—G. C. 8.
71. Sus verrucosus Miill. & Schleg.
gd. 662, 793, 882. 2. 666, 667, 669. Tjilatjap.
3. 1687, 1694 (imm.), 1708, 1785, 1786. 9. 1688, 1693
(imm.), 1782 (imm.). Pangandaran, Dirk de Vries Bay.
3. 1466. 2. 1467. Kalipoetjang, Tji-Tandoei River.
gd. 1149, 1305, 1366. 9. 1300. Tasikmalaja, Preanger.
392 ON FERTILIZED EGGS OF SEBA PYTHONS. | Apr. 6
“<«Babbi’ (Malay). ‘Tjelleng’ (Javanese). ‘ Bagong’
(Soendanese). ‘ Bagong-gadoong’ (Soendanese). ‘ Bagong-wraha.’
“Taking the place of Sus vittatws in the inland mountainous
districts of Java, but also occasionally occurring with it near
the coast.” —G. C.S.
. TRAGULUS FOCALINUS Miller.
1315. Batavia.
@. 1238. Tasikmalaja, Preanger.
@. 621, 683. Tyilatjap.
3. 1606, INGE OOS UTAORAS IECADS IS Tig IS Teeth pss,
2. 1646, 1667, 1676, 1696, 1697, 1700, 1701, 1704, 1705, 1713,
AA. Pangandaran, Dirk ds Wi 1es Bay.
“<Kantyil’ (Malay). ‘ Pootjang’ (Soendanese).
‘“‘Plentiful in dense forests, particularly in thinly populated
districts.
“T could get no information of more than one species of
Tragulus on the mainland of Java.”—G. C. 8.
73. Muntiacus MuNTIAK Zimm.
$6. 1771. @. 1732. Pangandaran, Dirk de Vries Bay.
““¢ Kidang’ (Javanese). ‘Muntjac’ (Malay, Soendanese).
“The uncultivated parts of Java, particularly in hilly
situations.” —G, C.S.
74. MANISs JAVANICA Desm.
6. 210, 211. Buitenzorg.
3. 1005. 'Tasikmalaja, Preanger.
6. 1306. Batavia.
©. 1691. Pangandaran, Dirk de Vries Bay.
“<¢ Poising’ (Soendanese). ‘ Tringilling’ (J ovens ae
“ Fairly plentiful, chiefly in forest country.”—G. C. §
April 6, 1909.
FREDERICK GILLETT, Esq., Vice-President, in the Chair.
The Secretary exhibited, on behalf of Mr. George Jennison,
some fertilized eggs from a pair of Seba Pythons in the Belle Vue
Zoological; Gardens, Manchester, and read the following account
of them :
Sythe Belle Vue snake-cage is 75 feet long by 10 by 12, forming
an annexe toa greenhouse and et (except the back) entirely
of glass. A minimum temperature of 80° is aimed at, and an
inner case, in which sat the Seba Python mentioned below, is
1909.) ON AN OX-BONE GNAWED BY RODENTS. 393
provided for retreat from extreme cold. Numerous hanging
baskets of growing plants and a wired-off portion which is one
mass of tropical vegetation beautify the cage, and make an
atmosphere and surroundings congenial to the serpents. These
things have had, I believe, great influence in the production of
eggs. Only one ‘batch was laid during 10 years in our old snake-
Cages, whereas, beginning April 1904, there have been six lots of
eggs in the new cage, which was first occupied in the summer of
1903.
“ Hitherto, with perhaps one exception, the eggs have not been
fertilized, but only once has the snake failed to incubate them for
a considerable period. And on one occasion a Python Molurus
permitted herself and eggs to be moved from one situation to
another without decreasing her ardour.
“The sitting of a Seba Python, which commenced Dec. 21st
and terminated at the 4th of February this year, would probably
have been successful had she not been lying on an air-grating, where
the temperature would approximate to 120°. The eggs, which
were perfectly white, through the whole period were pressed and
flattened out of spherical shape until they closely resembled a plaque
of white excreta. The reptile sat upon them very assiduously,
but left the nest for about 10 minutes morning and evening to
drink, and for about half an hour every fortnight when feeding,
which she did regularly on dead rabbits ‘during the whole period.
“ During the period of incubation the Python grew more vicious,
turning upon the keeper several times and occasionally striking
at visitors.
“ Nevertheless, she in no way resented the presence of the dozen
or more snakes which crowded into the same case in the cold
January days, but which usually kept as far from her as possible
(about 5 5 feet), the male Python alone lying at her side.
“This pair of Seba Pythons was purchased from Mr. Lyster
Jameson Jast summer, and had been, I believe, on deposit at
Regent's Park for some time previously.
“Slough of female 15 feet long.”
Dr. R. T. Leiper, F.Z.S., exhibited a greatly distorted Elephant’s
tusk from the Congo, and a malformed canine tooth of a Hippo-
mus from Uganda, and stdted that the conditions most
probably had originated from mechanical injury.
Mr. E. T. Newton, F.B.S., F.Z.S8., exhibited 2 metatarsal bone
of an Ox, showing in a remarkable manner the marks of gna
by rodents, possibly Squirrels, Rats, or Mice, which he had found
in the woods near Cromer.
Pezoc. Zoou. Soc.—1909, No. XX VI. 26
394 MR. R. I. POCOCK ON A [ Apr. 6,
The following papers were read :-—
1. Description of a new Form of Ratel (Mellivora) from
Sierra Leone, with Notes upon the described African
Forms of this Genus. By R. I. Pocock, F.L.S., F.Z.S.,
Superintendent of the Gardens.
[ Received March 9, 1909.]
(Plate LX.)
The Society has recently received from Mrs. Bettington a Ratel
which not only came frem Sierra Leone, a locality whence the
genus Mellivora has never been previously recorded, but differs
markedly in coloration from all the examples that I have seen
alive and equally so from all the skins, with one exception, that
are contained in the collection of the British Museum.
I propose to describe it as a new species under the name
Mellivora signata.
MELLIVORA SIGNATA, sp. n. (Plate LXT.)
Forehead and fore part of the nape and sides of the neck quite
white, the white on the sides extending farther back than on the
upper surface of the neck. In the angle thus defined upon the
posterior part of the nape, as well as on the shoulder, the colour is
jron-grey, being a mixture of wholly black and wholly white
hairs and of black hairs with a pale subapical annulus, the black
predominating. Behind the shoulders the prevailing colour of
the back is black ticked or speckled with yellowish and brownish
grey. None of the hairs of this region appears to be wholly pale
as in all other Ratels, except JZ. cottont; but a considerable number
of them have a narrower or broader pale subapical annulus
imparting the speckled appearance above described. Over the
hind-quarters this speckling gradually dies away ; there 1s, indeed,
a gradual but marked decrease in the amount of speckling from
the fore part to the hind part of the back, the speckling of the
fore part blending with the iron-grey hue of the shoulders and
the latter blending to a certain extent with the white of the head
and neck, though the angular line of demarcation is very visible.
Tail all black. Legs, muzzle, and under parts black, as in all
Ratels. Hairs on the sides of the neck showing a tendency to
grow forwards and inwards from a whorl on the shoulders, thus
forming a weak nuchal crest.
Loc. About 100 miles inland of Sierra Leone.
It was stated above that, with one exception, this specimen,
which is not yet adult, differs markedly from all the skins of
Mellivora in the British Museum. This exception is a piece of
the skin of the hind-quarters, flat and native-prepared, which
was brought from West Africa by Winwood Reade, together
with other material for which no locality was in any case,
12 A'S) NS) OS) le) LPL
Glebe Presslmp.
j
fe erep ark Lith.
i\
IMU EIL EWI) IRUAN tS) IMEINUAN IDEN,
1909. ] NEW RATEL FROM SIERRA LEONE. 395
I believe, certainly known. But the vessel in which Reade went to
the Gaboon stopped at the Gambia and Sierra Leone*, and it is
possible that the skin in question came from the latter place.
In any case it seems to belong to the same kind of Ratel as the
one I have here described as new, though possibly subspecifically
distinct from it. It is markedly dark in colour, being dark brown
speckled with dirty yellow. Some of the hairs are dirty yellow
throughout ; others have a pale annulus of that tint.
Although the marked contrast between the colouring of the
head and that of the body distinguishes J. signata at a glance
from its congeners known to me, a specimen described by Pousar-
gues from the French Congo appears to show a decided approach
to it, despite Pousargues’ remark that he could establish no differ-
ence between this specimen and examples from East and South
Africa. He states, however, that the back bears a median iron-
grey longitudinal zone, which extends from the middle of the
upper side of the neck to the tail and is limited in front by a
pure white “caloette” on the nape of the neck and head. It is
obvious, however, that this Congoese specimen differs entirely
from the one here described in the uniform iron-grey hue of the
back, shoulders, and hind quarters.
Mrs. Bettington, who brought this Ratel from Sierra Leone,
gave me the following information about it. According to native
reports the species is nocturnal and lives to a great extent upon
fish, which it catches with its paws at the edge of streams, and
the specimen above described preferred fish to meat during the
time she had it in captivity. It also showed a liking for bread
in preference to meat. She kept it in a yard with some fowls
and it never made any attempt to kill them. Thinking from
this that it could be trusted with other living animals, she
allowed it access one day to some puppies, but these it proceeded
to devour without delay. When she put it with an Irish terrier
the two merely sniffed each other and parted, although the terrier
would on ordinary occasions attack, and if possible kill, any wild
animals—such as civets—it came across. ‘This incident has an
interesting bearing upon the theory that the Ratel is protected
by its scent from attacks by large carnivora, a protection with
which is associated what is believed to be the warning characters
of its coloration T.
Unfortunately the animal died from a form of “ distemper,”
probably contracted on the voyage, a few weeks after being
deposited in the Gardens. Its anal stink-glands resembled in
position those of Skunks (Mephitis) although smaller. There was
a pair of them, each about the size of a thrush’s egg, opening
upon a papilla or nipple, just within the anal orifice, and lying
obliquely backwards deep in the surrounding tissue. The
* See ‘Adventures in the Forest and the Field,’ by The Old Shekarry (H. A.
Leveson), pp. 160-278.
+ Pocock, P. Z. 8. 1908, pp. 949-951,
26*
396 MR. R. I. POCOCK ON A [ Apr. 6,
external or adanal portion of the gland consisted of spongy tissue
traversed by a narrow passage leading to the aperture on the
papilla. ‘This no doubt was the secreting area. ‘The internal or
adoral portion, on the contrary, formed a hollow semi-oval space,
which apparently acts as a reservoir for the storage of the
ejectable fluid.
The discovery of this form of J/ellivora in the Sierra Leone
district considerably extends the known geographical range of
the genus, which had not been previously recorded to the west
of Lake Chad. It was also known to occur in the district between
the Niger and the Congo, in the Ituri Forest, and in East and
South Africa from Abyssinia and Somaliland to Cape Colony
and Angola.
The Kast and South African Ratels have always been referred
to the same species, JZ. ratel Sparrm., of which J/. capensis Schy.,
M. mellivora G. Cuv., and IM. typicus A. Sm. are absolute
synonyms. There does not seem to have been sufficient material
collected in different parts of East Africa to show whether these
animals have been differentiated into subspecies distinguishable
from typical J. ratel or not; but there is no doubt that specimens,
from Suakin for example, are very like those from Cape Colony.
Judging, nevertheless, from material in the British Museum, it
seems that South African specimens have on an average more
black hairs in the dorsal pelage and the white marginal line
more strongly pronounced than in North-east African examples.
Three distinct species, however, have been described from
West African material, namely :—
1. Mellivora leuconota P. L. Sclater (P. Z. 8. 1867, p. 98,
pl. viil., and 1871, p. 232). Based upon a young specimen from
West Africa which was separated from J/. ratel on account of the
nearly uniform whiteness of its upper side. When adult, the
animal exhibited more grey on the posterior region of the back.
This change in colour induced Dr. Sclater to abandon the species,
a conclusion in which he was followed by Pousargues (Ann. Sci.
Nat. iii. p. 275, 1896) and by W. L. Sclater (Fauna of 8. Afr.,
Mamm. i. p. 110, 1900). P. L. Sclater, however, remarked that
even the adult animal had the crown white; and his description
of the dorsal surface does not fit that of examples of the typical
8. African J/. ratel in the British Museum, which have the head
and back dark iron-grey, with a large percentage of black hairs
mixed with the white.
W. L. Sclater’s description of specimens in the Museum at Cape
Town, of which he says “ general colour above from forehead to
base of tail greyish brown, becoming darker posteriorly,” further
shows that the head is not, so far as is known, white in the
typical form. This is also apparently true of an example from
Suakin, which presumably formed the basis of the description and
figure of this animal in Anderson’s and de Winton’s ‘ Mammals
of Egypt,’ p. 246, 1902.
1909. ] NEW RATEL FROM SIERRA LEONE. 397
Pousargues, moreover, as above stated, described a specimen
from the French Congo as having the fore part of the neck and
the head white and the posterior part of the neck and the back
iron-grey. His description indicates a decided contrast between
the tints of the head and back. Finally, there is in the British
Museum a skin from W. Africa, sent home by Mr. G. L. Bates,
probably from the Cameroons or the Benito River, which is
uniformly yellow-white from the forehead to the root of the tail,
thus resembling the first published description of J/. lewconota,
These facts, coupled with the lack of evidence that the young of
typical I. ratel is whiter than the adult, induce me to believe
that the name lewconota should be preserved as a subspecific title
for, at all events, one of the Ratels, if there be more than one,
from the Congoese area.
2. M. cottoni Lydekker (P. Z. 8. 1906, i. p. 112, pl. vii.), from
the Ituri Forest. Quite distinct from all the previously described
Ratels in the absence of white on the dorsal area, the whole
animal being black. As Mr. Lydekker suggested, this may be a
melanotic sport; but there is no evidence of this up to the present
time, only one example being known from the locality.
Of the Ratels that I have seen, the one that comes nearest to
MM. cottont in colour is an example which was purchased by the
Society more than nineteen years ago from Cross, of Liverpool,
and is still living in the Gardens. Its locality is unknown. It
was entered in our books as ‘“‘ Mellivora capensis,” but the specific
name was subsequently altered to ‘ indica”—I suppose because
there is no whitish band defining the greyish hue of the dorsal
surface from the black of the “underside, this being formerly
regarded as the one distinguishing feature between the African
and Indian forms. It is an ‘unusually dark animal, darker, in fact,
upen the head and dorsal surface than any that I have seen
except J/. cottoni. These areas are very dark grey, being black
merely sprinkled with grey. One of the keepers, however, “Hoare,
who has known the animal for nearly twenty years, tells me that
he thinks it was a good deal greyer when first purchased as a
young animal. Another keeper, Dixon, on the contrary, who has
looked after it for fifteen years, says that it is the same colour
now as when he first took charge of it. However that may be,
there is no doubt that at the present time the animal is inter-
mediate in colour between typical Jf. ratel and WM. cottoni. I
cannot determine it more accurately. It may be that the dark
hue of the dorsal surface is a sign of age. In favour of this view
may be alleged the increase in the dark hue of the dorsal
area with advancing years in the type of J. leuconota; and
the fact that Blanford describes MM, indica as light grey or
whitish grey above, whereas in two Indian specimens in the
British Museum, both young animals, one from Nepal and the
other from Rajpootana, the dorsal area is almost wholly white.
If such a change typically takes place in both African and Indian
398 ON A NEW RATEL FROM SIERRA LEONE. [ Apr. 6
Ratels, the fact must be reckoned with in our estimate of the
characters of JZ. cottoni, and renders all the more remarkable the
very dark coloration of the back in the type of JZ. signata, a quite
young animal.
3. M. concisa Thomas & Wroughton (Ann. & Mag. Nat. Hist.
(7) xix. p. 376, 1907), from Yo, Lake Chad. Separated as a
species from J/. sel because it is a little smaller and because
there is a narrow median longitudinal black area extending from
the posterior end of the lumbar region, over the sacral region to
the root of the tail, which has no white hairs. The rest of the
upper surface is iron-grey, the head being only a little lighter
than the back, and there is no distinct white marginal stripe.
I find it impossible to give these characters more than subspecific
value. Indications of darkening of the sacral region may be seen
in skins from E. Africa scarcely separable from “typical M. ratel ;
and askin in the British Museum sent by Capt. Flower from
Khartoum presents nearly the same peculiarities in the coloration
of the hind-quarters as does the Lake Chad specimen. As for the
absence of the white hair from the root of the tail, no great
importance, it seems, can be attached to that character, for of two
skins in the British Museum, ticketed “‘ Somaliland, R. M.
Hawker,” one has no white hairs on the base of the tail, whereas
in the other the white hairs extend about two inches down that
organ. Only one example of J/. concisa was procured.
I subjoin a table of the named forms of African Ratels, leaving
for the present unsettled the question, not a very important one
in the present instance, as to the exact systematic rank that
should be assigned to them :—
a. No white hairs upon the head or back..................... cottoni, Ituri Forest.
a’. Head white or iron-grey ; a varying amount of grey
or dirty white or white on the back.
6. Head and fore part of the neck, especially at the
side, white and sharply defined from the vest of
the neck and back, on which the black is markedly
dominant, very few of the hairs being wholly
dirty w hite, this pale tint being confined to a sub-
apical annulus on some of the black hairs ............ signata, Sierra Leone.
bo’. Head and fore part of the neck sometimes w aintes
but never apparently very sharply defined by their
tint from that of the posterior part of the neck
and the body, which are either wholly white or
iron-grey, due to a mixture of hairs black or white
throughout their length.
e. Head white, back white (? young) or iron-grey ... lewconota, Congo.
c’. Head not white, iron-grey like the back though
a little paler.
d. No distinct marginal white band; sacral region
mesially black, no white on root of tail ...... concisa, L. Chad.
d’, A distinct marginal white band; hind-quarters
uniformly iron-grey at least to "the root of the
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1909.] ON A PROTOZOAN BLOOD-PARASITE OF THE SEA-TROUT. 399
2. Notes on an Ichthyosporidian causing a I’atal Disease in
Sea-Trout. By Muriet Ropertson, Carnegie Research
Fellow *.
[Received March 1, 1909.]
(Plates LXII—LXIV.7)
In July 1907 Dr. Turnbull, of the London Hospital Medical
School, while fishing on the River Ewe in Ross-shire, observed
that a number of sea-trout were affected with a disease which cul-
minated in the death of the fish. Externally they showed nothing
abnormal, but on opening them he found that the heart, liver,
spleen, and pyloric ceca showed minute sand-like granules.
Dr. Turnbull’s description of a typical infection reads as
follows :—
“The gills were healthy and showed no adherent parasites.
“The heart: the ventricle was pale, rough and sandy, and
minute white granules could be seen. Similar granules were on
the auricles and sinus venosus.
“ The liver was of a yellow colour and showed minute, yellowish-
white, slightly raised, rounded granules on the surface and in
section.
“‘Tn the spleen very few granules could be detected.
“ The alimentary tract was empty and no duodenal tape-worms
were present.
“The brain, on removal, showed no abnormalities.
“¢The ovaries showed no granules.”
Some fish, however, showed an even more complete infection
than this; thus in some cases the pyloric ceca, the gills, the
submucosa of the csophagus, the testicles, and the parietal
muscles beneath the peritoneum all showed the characteristic
sanded appearance.
Dr. Turnbull observed that the diseased sea-trout were all
fresh-run fish and were generally of about 4 lbs. weight and over,
although one infected fish was got which weighed 2 lbs.
The River Ewe runs from “‘ Loch Maree” into the sea-loch
“Loch Ewe” at “ Poolewe.” It is a short river, only about a
mile and half long.
The various internal organs of some of the infected fish were
preserved in methylated spirit, this bemg the only preserving-
fluid obtainable. Sections were made which were sent along with
Dr. Turnbull’s notes to Prof. Minchin at the Lister Institute.
The disease proved to be caused by a Protozoan parasite belonging
to the genus Jchthyosporidium.
In January 1909 Prof. Minchin handed over to me the
material together with his own and Dr. Turnbull’s notes, and it
* Communicated by Prof. E. A. Mrncurn, M.A., V.P.Z.S.
+ For explanation of the Plates see page 402.
400 MISS M. ROBERTSON ON A PROTOZOAN [Apr. 6
is from these that this account of the parasite has to a large
extent been drawn.
The genus Jcehkthyosporidium belongs to the family of the
Bertramide and to the order of the Haplosporidia. It was
created by Caullery and Mesnil in 1905 for the two parasites
Ichthyosporidium gasterophilum and TIchthyosporidium phymo-
genes *.
Ichthyosporidium gasterophilwm, which is closely allied to if not
identical with the parasite in the sea-trout, is found in Motella
mustela and Liparis vulgaris.
These are both shore-living fishes and inhabit the Fucus-belt.
Caullery and Mesnil note that J/otella mustela (the rockling) was
most frequently infected and that the infected fish were all above
a certain size. The parasite was found only in the pyloric ceca
and the ducts of the glands of the stomach. The authors do not
mention if the presence of the /ehthyosporidium seriously affected
the health of the fish.
The Jehthyosporidium of the sea-trout can easily be recognised
in sections of the tissues as large spherical or elongated bodies
(figs. 1-8). The material only shows the trophical stages. of the
parasite and the fixation is not quite without reproach.
The parasites are multinucleate organisms, most often spherical
in shape and possessing a well-marked outer envelope. They
range from large individuals with many hundred nuclei, measuring
120 mm. in diameter, to small forms with a thin envelope and few
nuclei. Individuals with only two nuclei are to be found and
some apparently with a single nucleus, but it is difficult to be
quite certain of these. They are very rare, and in section-
material of this type an error in such a point is very easily
made.
The envelope is secreted by the parasite; the inner part is
smooth and structureless except for occasional striations (figs. 6
& 7). The outer part of the envelope is often slightly crinkled.
The shape of the organism is open to very considerable variation ;
oval, vermiform, and even irregularly branched forms may be
found. These are for the most part individuals which have
quitted their original envelope and grown out into the tissues.
They seem to secrete a secondary envelope, which is generally
rather thin; moreover, this may not extend over the whole
animal.
Sometimes these large irregular specimens seem to be breaking
up into several daughter individuals by the very simple process
of plasmotomy. The products of this process are of very varying
sizes. It is a very casual method and does not involve nuclear
division—it is, in fact, merely the breaking off of a mass of nu-
cleated protoplasm.
* Caullery and Mesnil, C. R. Soe. Biol. Paris, lviii. (1905) pp. 640-642.
1909. ] BLOOD-PARASITE OF THE SEA-TROUT. 401
It is impossible to say exactly what brings about the exit from
the envelope. It is not a question of size, as quite small specimens
may be found fixed at what appears to be the moment of
escape.
The nuclei in Ichthyosporidium are, relatively speaking, very
small and very numerous. They consist each of a single compact
mass of chromatin surrounded by a clear space which is bounded
by a delicate sharply-defined membrane. Very fine rays pass
from the mass of chromatin to the membrane. The mass of
chromatin or karyosome is dense and compact, and shows no
structure, no matter what the stain used. I[ron-hematoxylin,
Delafield’s hematoxylin, cavm-alum and picronigrosin, picro-
carmine and picronigrosin, and Twort’s neutral red and light
green stain were all tried and all gave very fair results. Iron-
hematoxylin and Twort’s stain, however, gave the best pictures.
The rays passing from the karyosome to the outer membrane are
exceedingly fine and stain rather faintly with the chromatin
stains. The membrane takes up the chromatin colours faintly when
stained with iron-hematoxylin, Delafield’s hematoxylin, &c., but
takes up the green of the Twort’s stain—that is to say, it does
not take the chromatin stain of this combination. This is a
rather interestmg point, as it indicates that whatever its nature
it is not the mere condensation of chromatin which generally
constitutes the so-called “ membrane” in the Protozoa. It may
be noted that the red colour in Twort’s stain, when properly
applied, appears to be a pretty good test for chromatin.
No stages showing nuclear division were observed nor was
there any indication of spore-formation.
As has already been stated, the parasite invades practically all
the organs of the trout, but seems always to be found in the
greatest numbers in the muscles of the heart.
The reaction on the part of the host in the way of forming
connective-tissue cysts round the parasites is comparatively slight.
In the liver and spleen, it is true, cysts composed of many layers
of connective-tissue cells are formed, especially in the case of the
larger parasites; but in other parts of the body, such as the heart-
muscles, the cysts consist only of two or three thin layers of cells,
and may sometimes not be formed at all. The envelope secreted
by the parasite in these cases lies in direct contact with the
striped muscle-fibres. This relatively slight development of
connective tissue may have a certain importance in explaining
the fatal nature of the disease.
Three years ago, in January 1906, I came across an Ichthyo-
sporidium* identical, as far as I can see, with the one found in the
trout, in a smal] flounder (Plewronectes flesus).
The fish was also infected with a trypanosome, which was the
parasite I was in search of, and I unfortunately did not examine
% Proc. Roy. Phys. Soc. of Edinburgh, vol. xvii. No. 5, 1908.
402 ON A PROTOZOAN BLOOD PARASITE OF THE SEA-rROUT. { Apr. 6,
the heart. The liver and the submucosa of the stomach and the
intestine were very strongly infected, far more so than any of the
organs from the trout. This material gave a more complete
picture of the trophic life of the parasite. Such points as the
multiplication of the nuclei, the exit from the envelope, and the
breaking up of the large individuals into numerous small naked
bodies with few nuclei were clearly illustrated in the sections.
There were, however, nosigns of sporogony. In the flounder very
large connective-tissue cysts were formed both in the liver and in
the alimentary canal. The only difference between the infection
in the sea-trout and that in the flounder is in the greater
development of connective tissue on the part of the latter host.
The flounder in question must ultimately, I should think, have
succumbed to the disease, as the liver was in a very pathological
condition, but it had been living for some months in captivity
in the tanks of the Millport Marine Station and showed no
external sign of ill-health.
From the material at my disposal only a very incomplete
account of the Ichthyosporidium can be given. It is to be hoped
that this form, which is interesting both from a purely scientific
as well as an economic point of view, will receive further attention
from Protozoologists and will be studied under more favourable
conditions.
Since writing the above paper another case of the occurrence of
this disease has come under my notice in haddocks from Aberdeen.
A fishmonger in Glasgow, while cutting the fish into fillets,
observed minute yellowish white granules in the tissues. He
sent the fillets to Prof. Graham Kerr, who forwarded them to
me at the Lister Institute. It was found that the appearance
was due to the presence of an immense number of an Ichthyo-
sporidian apparently identical with the one described in this
paper.
EXPLANATION OF PLATES LXII-LXIV.
Ichthyosporidians of Sea-Trout.
Fig. 1. Magn. of 35 diam. Showing general appearance of the parasites in section
of the heart.
Magn. of 100 diam. General view of the parasites, showing different shapes
assumed.
Magn. of 250 diam. Single large parasite, showing the envelope.
Magn. of 250 diam. Single large parasite; the ‘Shrinkage is due to the
fixation, as also the appearance of the nuclei at one side. The envelope
shows clearly.
5. Magn. of 500 diam. Shows thin layer of connective tissue round the
arasite; envelope of the Ichthyosporidiwn and the nuclei are both shown.
Figs. 6 & 7. Magn. of 1000 diam. These two ficures show the nuclei; the fine rays
passing from the karyosome to the membrane can be seen.
Fig. 8. Magn of 1000 diam. Shows the nuclei distorted in the process of fixing.
9. Magn. of 750 diam. Gives another appearance caused by fixation.
bo
These figures are from untouched photographs kindly executed by Dr. Reid.
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3.SALARIAS CAUDOFASCIATUS. 4S NATALIS. 5.S.MELANOSOMA.
6.CIRRHITES MURRAYI.
1909.] ON FISHES FROM CHRISTMAS ISLAND. 403
3. A Collection of Fishes made by Dr. C. W. Andrews,
F.R.S., at Christmas Island. By C. Tare Reean,
M. As. EAS.
[Received March 4, 1909. |
(Plates LXV. & LXVI.*)
The fishes obtained by Dr. Andrews at Christmas Island are
of considerable interest, as they include examples of seven species
which appear to be new to science.
1. Murana NEBULOSA Ahl.
2, PLESIOPS MELAS Bleek.
3. SPHHZRODON HETERODON Bleek.
4, PARUPENEUS ANDREWSII, sp. n. (Plate LXV.)
Depth of body a little more than the length of head, which is
33 in the length of the fish. Interorbital region very convex,
its width 32 in the length of head; diameter of eye 6 in the
length of head; lips thick ; barbelsa little more than 3 the length
of head; 25 gill-rakers on the lower part of the anterior arch.
31 scales in a longitudinal series; tubules of lateral line much
branched. Dorsal VIII, 9; second spine slender, flexible, not
stronger than the others; third longest, 3 the length of head;
last soft ray moderately produced. Anal I, 7, the spine very
small. Pelvics a little longer than the pectorals, 2 the length of
head. Caudal peduncle nearly twice as long as deep. Reddish,
with two broad blackish vertical bands below the bases of the
spinous and soft dorsal fins respectively; snout dark bluish ;
spinous dorsal blackish; soft dorsal blackish at the base and
with somewhat oblique stripes superiorly; anal with similar
stripes.
A single specimen, 270 mm. in total length. From allied
species with a somewhat similar coloration P. andrewsi is dis-
tinguished by having the second dorsal spine slender and flexible.
Dr. Pellegrin has kindly examined the specimens in the Paris
Museum described by Cuvier and Valenciennes as P. bifasciatus
and P. trifasciatus, all of which have a strong, pungent, second
dorsal spine tf, as have the specimens in the British Museum
referred to those species. P. bifasciatus, in most respects the
nearest to P. andrewsi, differs also in that the last ray of the
second dorsal is the shortest instead of the longest (Bleek. &
Pollen, Faun. Madagasc., Poiss. pl. xviii. fig. 2).
5. CHELMO LONGIROSTRIS Brouss.
* For explanation of the Plates see page 406.
+ This was clearly seen in Lacepéde’s type of P. bifasciatus.
404 MR, C, TATE REGAN ON FISHES [ Apr. 6,
6. CIRRHITES MURRAYI, sp. n. (Plate LXVI. fig. 6.)
Depth of body 23 to 3 in the length, length of head 3} to 33.
Snout as long as or longer than eye, the diameter of which is
34 to 4in the length of head and about twice the interorbital
width. Maxillary extending to below anterior part or middle
of eye; anterior nostril with a fringed tentacle; cheek with
4 series of scales. 44 to 46 scales in a longitudinal series.
Dorsal X 12; fifth and sixth spines longest, 4 the length of head ;
interspinous membranes deeply notched, produced into a penicillated
lobe behind each spine; first soft ray the longest, about ? the
length of head. Anal III 6; second spine strongest and longest,
2 the length of head. Pectoral with six simple rays. Caudal
subtruncate. Reddish, with about 6 blackish vertical bars, each
of which is usually broken up into 3 or 4 oblong spots; blackish
spots on head and on spinous dorsal fin.
Seven specimens, 64 to 78 mm. in total length; a number of
smaller examples are not included in the description. The species
is named after Sir John Murray, K.C.B., F.R.S.
I accept the family Cirrhitide as defined and limited by
Dr. Giinther, with the addition of Haplodactylus. This very
natural group is characterized externally by a number of features,
amongst which may be mentioned the unbranched lower rays of
the pectoral fins and the posterior position of the pelvic fins. Of
osteological characters the most notable are the form of the
cleithra, which have strong horizontal expansions anteriorly and
meet in a long symphysis inferiorly, and the attachment of the
well-developed epipleurals at or immediately above the insertion
of the ribs. A subocular shelf is present in Cirrhités and Chilo-
dactylus, absent in Chironemus, Latris, and Haplodactylus.
Parapophyses commence on the first vertebra in Latris, Haplo-
dactylus, and Chilodactylus, on the third or fourth in Chironemus,
and on the fifth or sixth in Cirrhites.
7. GLYPHIDODON soRDIDUS Riipp.
8. PLATYGLOSSUS HYRTELII Bleek.
9, PrrRois votirans Linn.
10. CARACANTHUS UNIPINNA Gray.
11. Gosius ALBOPUNCTATUS Cuv. & Val.
12. TRIPTERYGIUM ATROGULARE Giinth.
13. BLENNIUS NATIVITATIS, sp.n. (Plate LX VI. fig. 2.)
Depth of body nearly equal to length of head, about 5 in the
length of the fish. Snout vertical; maxillary extending to below
posterior edge of eye; a pair of posterior canines in the lower
1909. ] FROM CHRISTMAS ISLAND. 405
jaw; tentacles all simple, the nasal and nuchal pairs short, the
ocular pair longer ; no crest on the head. Dorsal XII 15; notch
rather deep ; last ray not connected with the caudal. Anal 18-20.
Caudal rounded or subtrunecate. Reddish, with about 8 darker
cross-bars, sometimes appearing only as a series of oblong blotches
along the middle of the sides ; fins pale.
Several specimens, measuring up to 45 mm. in total length.
14. BLENNIUS ATROCINCTUS, sp.n. (Plate LXVI. fig. 1.) |
Depth of body nearly equal to the length of head, about 5 in
the length of the fish. Snout vertical; maxillary extending to
below middle of eye; no canines; nasal and orbital tentacles
bifid ; no nuchal tentacles; no nuchal crest. Dorsal XIII 17;
first three rays rather wide apart; notch rather deep; last ray
not connected with the caudal. Anal 22. Caudal rounded or
subtruneate. Reddish, with 6 blackish cross-bars continued on
to the basal part of the dorsal fin ; anal blackish.
A single specimen, 27 mm. in total length.
15. SALARIAS HASSELTII Bleek.
16. SALARIAS CAUDOFASCIATUS, sp. n. (Plate LX VI. fig. 3.)
Allied to S. marmoratus Benn. Depth of body a little less
than the length of head, + the length of the fish. Snout vertical ;
maxillary extending to below posterior edge of eye; a pair of
posterior canines in the lower jaw; nasal and orbital tentacles
fringed; a pair of short, simple nuchal tentacles; no crest.
Dorsal XIT 15; anterior portion low; second portion elevated,
with the rays decreasing in length from the first; last ray not
joimed to the caudal. Anal 18. Caudal subtruncate. Reddish,
with scattered pale spots and with 6 dark cross-bars, each of
which is split into two; caudal with dark vertical bars; anal
dusky.
A single specimen, 55 mm. in total length.
17. SALARIAS NATALIS, sp. n. (Plate LXVI. fig. 4.)
Allied to S. unicolor Riipp. Depth of body equal to or less
than the length of head, 54 to 6 in the length of the fish. Snout
vertical ; maxillary extending to below posterior edge of eye or
beyond; a pair of posterior canines in the lower jaw; nasal
tentacles simple or fringed ; orbital tentacles rather long, fringed ;
no nuchal tentacles; an occipital crest. Dorsal XII—XIIT 17-18;
notch deep; last ray not joined to the caudal. Anal 20-22.
Caudal subtruncate or rounded. Head and body with 14 purplish
cross-bars arranged in pairs and usually split up to form 3 longi-
tudinal series of spots; fins dusky, the dorsal with oblique
stripes.
Eleven specimens, measuring up to 60 mm. in total length.
406 MR. H. H. DRUCE ON BUTTERFLIES [ Apr. 6,
18. SALARIAS MELANOSOMA, sp. n. (Plate LXVI. fig. 5.)
Allied to S. anomalus Regan (Journ. Bombay Nat. Hist. Soc.
xvi. 1905, p. 327, pl. ii. fig. 4), but with the nasal tentacles double
or even triple, one or sometimes two shorter tentacles originating
at the base of the longer one. Dorsal XII 17, the spines not
produced, the longest not more than 4 the depth of the body.
Anal 20. Caudal truneate or slightly trilobed. Blackish or dark
brownish ; middle rays of caudal dusky.
Six specimens, measuring up to 60 mm. in total length.
The presence of posterior canines in the lower jaw led me to
re-examine S. anomalus and I find they are present, but often
hidden by a fleshy fold.
19. BroruLA MULTIBARBATA Schleg.
20. BALISTES RECTANGULUS Bl. Schn.
21. Diopon HystRix Linn.
EXPLANATION OF THE PLATES.
Pratt LXV.
Parupeneus andrewwsii.
Pratt LXVI.
. Blennius atrocinctus.
5 nativitatis.
. Salarias caudofasciatus.
Ss natalis.
: 3 melanosoma.
. Cirrhites murrayi.
Fig.
DOV Cot
4, On some New and Little-known Hesperiide from Tropical
West Africa. By Haminron H. Drucs, F.LS.,
Ze:
{ Received March 16, 1909.]
(Plate LX VIL.*)
Amongst a collection of Lepidoptera obtained in the Cameroons,
T have been able to distinguish 74 species of Hesperiide, including
those described in this paper. Professor Aurivillius has recorded
62 in his list of Butterflies from the Cameroons (Ent. Tidsk.
1896, pp. 279-291), and the following species are not referred to
by him, 32 in all, making about 100.
Types of new species in Mus. Druce,
* For explanation of the Plate see p. 413.
12, ZS LOS, 1k, WON
Y
Horace Knight del.et lith. West, Newman chr.
NEW AND LITTLE KNOWN W. AFRICAN HESPERIIDA:.
1909. | FROM TROPICAL WEST AFRICA. 407
Celenorhinus nigropunctata Parnara fatuellus Hopf.
B.-Baker. 3 noctula, sp. D.
Ortholexis hollandi, n. n. » palocampta, sp. 0.
Sarangesa brigidella Plotz. 53 flavifasciola, sp. n.
- bouviert Mabille. Pardaleodes incerta Snellen.
i. perpaupera Holland. | bule Holland.
Abantis elegantula Mabille. 3 astrape Holland.
» efulensis Holland. % ligora Hew.
PA WR easy, 5 vibius Hew.
» lucretia, sp. 0. _ Ceratrichia dinidiata Heron.
Hesperia spio Linn. | Andronymus leander Plotz.
Acleros olaus Plotz. Cenides kanguensis Holland.
» sparsum, sp. 0D. BF leonora Plotz.
Rhopalocampta bize Linn. » dacena Hew.
a unicolor Mabille 5 lacida Hew.
Ee hanno Plotz. ,» zaremba Holland (nec
Osmodes adon Mabille. Plotz).
Parnara arela Mabille. As laterculus Holland.
3 subochracea Holland. » terda Moschl. 2
: leucophea Holland. » cerynuca Hew.
es borbonica Boisd.
CELENORHINUS NigRoPUNCTATA. (Plate LXVII. fig. 11.)
Celenorhinus nigropunctata Bethune-Baker, Ann. & Mag. Nat.
Hist. ser. 8, vol. ii. p. 478 (1998).
I have figured a specimen from Bitje, Ja River, Cameroons,
which is identical with Mr. Bethune-Baker’s type from Makala,
Congo Free State. It is much like C. rutilans *, but the fore
wings are shorter and differently marked.
ORTHOLEXIS MELICHROPTERA. (Plate LX VII. fig. 1.)
Ortholexis melichroptera Karsch, Ent. Nach. 1895, p. 320, 3;
Aurivillius, Ent. Tidskr. 1896, p. 288, 2.
Both Herr Karsch and Professor Aurivillius record this species
from the Cameroons. I have figured a female obtained at Bitje,
Ja River, from the same district, by Mr. G. L. Bates, in the dry
season (2000 feet). It is apparently unknown to M. Mabille, as
he does not mention it in his ‘“‘ Hesperide” in ‘Genera Insec-
torum.’? Dr. Holland’s genus Acallopistes (P. Z.8. 1896, p. 95)
seems very nearly related, but the figure and description of the
antenne do not agree. Unfortunately we do not possess either
of the species he places in it; neither are they in the British
Museum.
ORTHOLEXIS HOLLANDI, nom. nov.
Katreus johnston Holland, Ent. News, v. pl. 1. fig. 8 (nec
Butler).
* Pardaleodes rutilans Mab. Bull. Soc. Zool. France, 1877, p. 235.
408 MR. H. H. DRUCE ON BUTTERFLIES [Apr. 6,
Hab. Bitje, Ja River, Cameroons, 2000 feet (G. ZL. Bates).
Dry season. Upper Kasai District, Congo Free State (C. Land-
beck).
Professor Aurivillius seems to have been the first to notice that
Dr. Holland had “ initiated a remarkable error in respect to this
form” [KEnt. Tidskr. 1896, p. 288] in figuring it as Katreus john-
stont Butler. It is, however, possible that Dr. Holland con-
sidered it to be the male of that species, but he gave no descriptive
account of the insect. The type of Choristoneura Jjohnstont
Butler is a male, and is slightly darker than females in the
British Museum, and has a small clear shining patch near the
base of the fore wing below, but otherwise the sexes are similar.
We have one male from Bitje and three males from the Congo,
but I have not seen the female.
In the form of the antenne and the palpi, and also in the
venation, this insect agrees well with Ovtholexis melichroptera,
of which I was inclined to think it was the male. The short
subapical fascia is semihyaline white, aml the upper surface of
both wings has a slight flush of blue. The cilia of the hind wing
on both surfaces is whitish, that of the fore wing being con-
colorous with the wings. The palpi and legs are br ioht yellow.
M. Mabille places the genera Aatreus and Chor ‘istoneura widely
apart in his “ Hesperidee” (1904), although au Heron, as long
ago as 1898 (Ann. & Mag. Nat. Hist. (7 ya p. 182), has written
that they are synonymous.
ABANTIS JA, sp.n. (Plate LX VII. fig. 2.)
Male. Upper side allied to A. elegantula Mabille; fore wing
coloured as in that species, and with the four discal and three
subapical hyaline spots arranged likewise. Hind wing with the
outer marginal third paler than median area, and with the basal
area whitish hyaline crossed by the black nervules. Cilia dark
glistening brown. Under side: both wings greyish buff, palest
along aed inner margin of fore wing and [paternal fold alk hind
wing, with black veins and hyaline spotsas above. Thorax black,
with ‘the convergent tufts of hair brick-red. Abdomen greyish,
with a central dark line above and broadly pale buff’ below.
Venter dark brown. Antenne black. Palpi black above with a
white spot ; yee. below, terminal joint black. Legs yellow.
Expanse 1,5 inch.
Hab. Bitje, ae River, Cameroons, 2000 feet (G. Z. Bates). Dry
season.
Distinguished at once from A. elegantula* by the hyaline
basal area on the hind wing and by the absence of the yellow
basal suffusion on the hind wing above and the unicolorous hind
wing below, the basal and discal areas of A. elegantula being
white on that wing.
* Abantis elegantula Mab. Ann. Soc. Ent. France, 1890, p. 32; Novit. Lepid.
p. 23, pl. iii. fig. 6 (1891).
1909, | FROM TROPICAL WEST AFRICA, 409
Mr. Bates captured specimens of A, elegantula in both the dry
and the wet seasons, but they show no difference in coloration or
size.
ABANTIS LUCRETIA, sp.n, (Plate LX VII. fig. 3.)
Male. Upper side: tore wing dull blackish brown, with three
subapical and a discal series of four white spots much as in
A, elegantula, the upper cell-spot being, however, very minute,
almost obsolete. ‘lhe nervules are darker than the ground-colour,
At the base of the wing, placed between the median and sub-
median nervures, is a dull red patch merging into the ground-
colour, Hind wing blackish brown, crossed from the costa, which
is orange, to the abdominal fold, which is black, by a broad pure
white band, which is sharply defined basally, but which has its
outer edge irregular on account of the black nervules reaching
beyond its border. An anteciliary row of white spots towards
the anal angle. Under side as above but paler; fore wing
without the dull red patch at the base. Hind wing with the
costa pale brown, a black dash at the origin of the subcostal
nervure, and the anal area broadly black. White spots on the
collar and the end of the patagia. Abdomen white, with a central
black streak above and a black streak on either side below.
Venter blackish. Antenne black; legs yellow; palpi black
above, yellow below, terminal joint black. Cilia above and below
concolorous with wings except towards anal angle, where it is
tipped with glistening white.
Expanse 12 inch.
Hab, Bitje, Ja River, Cameroons, 2000 feet (G. LZ. Bates).
Wet season.
T have named this insect, which is not close to any described
in the genus, after Pseudacrwa lucretia Cr.*, to which it bears a
close colour resemblance, and of which specimens were contained
in the same collection.
ABANTIS EFULENSIS.
Abantis efulensis Holland, P. Z.8. 1896, p. 21, pl, v. fig. 12, 3.
Male. Upper side: ground-colour paler than in male; white
areas of hind wing almost obsolete, being reduced toa short, streak
at end of cell and a small patch at anal angle. Under side as
male, but slightly paler. Abdomen, which has the base of each
segment black, is yellowish brown above and white below.
Hab. Bitje, Ja River, Cameroons, 2000 feet (G@. ZL. Lutes).
Dry season.
A male taken at the same time and place has the abdomen
white above and below, with the base of each segment black, in
this respect only not agreeing well with Dr. Holland’s figure,
which is doubtless taken from a stained specimen,
* Papilio lucretia Cramer, Pap. Ex. i. t. 45. figs. C, D (1779),
Proc. Zoot. Soc.—1909, No. XX VII. 27
410 MR. H. H. DRUCE ON BUTTERFLIES [Apr. 6,
ACLEROS SPARSUM, sp.n. (Plate LX VII. fig. 4.)
Male. Upper side: both wings uniform blackish brown, fore
wing without markings, hind wing with the anal angle and
outer margins almost to the apex irregularly pure white.
Under side: fore wing blackish brown, with the apex and outer
margin laved with white and grey, darkest along the costa.
Hind wing laved with grey and white, whitest towards the base
of the costa and at the anal angle. Thorax black above, white
below. Abdomen black above, white towards anus and white
below. Palpi: first jomt white, second joint white at the base
then buff interspersed with black, terminal joint black. Legs
black and white, spurs white. The antenne are black, with the
usual pure white spot just before the apex.
Expanse 1? inch.
Hab. Bitje, Ja River, Cameroons, 2000 feet (G@. LZ. Bates).
Dry season.
Described from two specimens,
This is a large species, and is perhaps nearest to A. lewcopyga
Mabille*, originally described from Madagascar, but found to also
inhabit W. Africa, but has less white on the hind wing above
and is differently marked below.
ACLEROS KASAI, sp.n. (Plate LX VII. fig. 5.)
Female. Upper side light brown with whitish hyaline spots.
Fore wing: three minute spots placed in a line from the costa, the
lower one being the largest, about halfway between the end of
the cell and the apex. Two spots in the cell at the end placed
one above the other; a spot between the two lower median
nervules and a smaller spot above and resting on the submedian
nervure about its middle. Cilia brown. Hind wing with a
distinct double spot forming a triangle towards the end of
the cell. Anal angle, outer margin, and cilia pure white.
Under side—Fore wing: costa rusty brown, discal area brown,
spots as on upper side, outer margin with a pale pinkish-brown
patch below the apex. Central inner marginal area broadly
whitish. Cilia pale brown. Hind wing laved with pinkish
brown, with darker shadings at the apex and along the inner
margin. An outer marginal row of darker brown shades. Cell-
spot as above, brown-edged. Anal angle white. Cilia white.
Abdomen brown above, with the ventral half pure white; white
below. Antenne brown, with the usual white spot on the club
just before the tip.
Expanse 12 inch.
Hab. Upper Kasai District, Congo Free State (P. Landbeck).
This is a large species, not very closely allied to any which I
can find described. The clear spots on the fore wing and the
distinct central spot on the hind wing at once distinguish it.
* Cyclopides leucopyga Mabille, Bull. Soc. Ent. France, 1877, p. 101.
1909.] FROM TROPICAL WEST AFRICA. 41}
GORGYRA AFIKPO, sp.u. (Plate LXVII. fig. 6.)
Male. Allied to G, abure Plotz*. Upper side: ground-colour.
blackish brown with whitish hyaline spots. Fore wing: a
subapical series of three minute dots, the innermost placed
further inwards, an elongated spot in the cell near the end, and
a smaller, more circular spot above it. A series of four discal
spots placed between the nervules nearest their origins, the
lower one resting on the submedian nervure about its middle.
Hind wing with a large ¥-shaped spot in the cell and two large
spots rather beyond the centre of the disc divided by the lower
median nervule. The patch of hairs on vein 16 is grey.
Under side: hyaline spots as above; fore wing with the apex
and outer margins washed with greyish. Hind wing grey, with
black spots arranged as in G. abure but much larger. Thorax
and abdomen concolorous with wings. Palpi pale straw-colour
below, black above, terminal joint black. Legs brown; fore
coxe clothed with thick straw-colour hairs. Antenne black.
Expanse 13 inch.
Hab. Afikpo, N. Nigeria (Reddick).
Unlike G. abure this insect bears but one pair of spurs on the
hind tibiz. In general appearance it seems to be much like
some species of the genus Gastrocheta Holland, viz. G. mabillei
Holland? and G. cybeutes Holland, but the palpi prove it to
belong to Gorgyra.
I have shown this insect to M. Mabille, who states it is quite
unknown to him.
PARNARA NOCTULA, sp.n. (Plate LXVII. fig. 7.)
Male. Upper side: both wings unicolorous dark bronze-brown
without markings. Cilia concolorous with wings. Under side as
above. Head, thorax, and abdomen dark brown; after part of
thorax and base of abdomen clothed with dark green hairs.
Legs dark brown. Palpi thickly clothed with black and white
hairs. Antenne black, tips chestnut-brown.
Expanse 13 inch.
Hab. Bitje, Ja River, Cameroons, 2000 feet (G. Z. Bates).
Dry season.
The total absence of markings should distinguish this from all
other described species of Parnara.
PARNARA PALOCAMPTA, sp.n. (Plate LX VIL. fig. 8.)
Female. Upper side: both wings uniform dark brown without
markings. Cilia of fore wing concolorous with wings; cilia of
hind wing from apex to anal angle whitish. Under side as
above. Head, thorax, abdomen, and legs blackish brown.
* Apaustus abure Plétz, S. e. Z. vol. xl. p. 359 (1879), MS. drawings pl. 736.
Holland, P. Z.S. 1896, pl. ii. fig. 18, as Katreus johnstonii Butler, error.
+ G. mabillei Holland, Ent. News, vol. v. p. 28, pl. i. figs. 15, 16 (1894).
ft G. cybeutes Holland, Ent. News, vol. v. p. 94, pl. iii. fig. 15 (1894).
Pine
412 MR. H, H, DRUCE ON BUTTERFLIES [Apr. 6,
Palpi thickly clothed with black and white hairs. Antenne
black, excepting tips and outer half of club, which are chestnut-
brown.
Expanse 1,4 inch.
Hab. Bitje, Ja River, Cameroons, 3000 feet (G@. ZL. Bates).
Dry season.
- This insect in coloration and shape is very much like a small
specimen of Rhopalocampta unicolor Mabille*, which we have
received from the same locality, but an examination of the
palpi proves that it is not congeneric with that species. The
fore wings are more elongated and the hind wings are less
produced posteriorly than im most species of Parnara. It seemed
possible that this might be the female of the previously described
Parnara noctula, but besides the white fringe on the hind wing
and the different shape of the wings the antenne present
considerable differences—they are shorter, the shaft is stouter,
and the club more graduated.
PARNARA FLAVIFASCIOLA, sp. n. (Plate LXVII. fig. 9.)
Male. Allied to P. tarace Mabillet. Fore wing dark bronze-
brown, with a central row of irregular white spots arranged as in
that species. Hind wing with the central yellow fascia broader
and without the yellow basal spot. Under side :—Fore wing:
ground-colour paler and with spots as above ; hind wing much
like that of P. statirides Holland ?, but yellow area rather less
extensive and base clouded with brown. Head black, with a
yellow central streak and yellow around the eyes. Thorax
brown. Abdomen brown above, yellow below, and with a yellow
band at the anus; venter brown. Legs brown with yellow
hairs. Palpi black above, yellow below, terminal joint black.
Antenne black, tips yellow.
Expanse 14 inch.
Hab. Bitje, Ja River, Cameroons, 2000 feet (G. LZ. Bates).
Dry season.
There is a specimen in the British Museum from Sierra Leone
from the Crowley Bequest.
PARNARA ARELA.
Hypoleucis arela Mabille, C. R. Soc. Ent. Belg. xxxv. 1891,
p. lxix.
Baoris arela Holland, P. Z.S. 1896, p. 68, pl. 11. fig. 20.
Ceratrichia paucipunctata Bethune-Baker, Ann. & Mag. Nat.
Hist. ser. 8, vol. ii. p. 481 (1908).
Mr. Bethune-Baker has shown me his type, which is without
doubt a male specimen of this common species with the antenne
wanting.
* Ismene unicolor Mabille, Ann. Soc. Ent. France, (5) vol. vii. p. xxix (1877).
+ Pamphila tarace Mabille, C. R. Soc. Ent. Belg. vol. xxxv. p. elxxix (1891);
Novitates Lepid. p. 114, pl. xvi. fig. 1 (1893). ‘
t Baoris statirides Holland, P.Z.S. 1896, p. 69, pl. v. fig. 6.
1909. | FROM TROPICAL WEST AFRICA. — 413
PARDALEODES BULE.
Pardaleodes bule Holland, P. Z.S. 1896, p. 76, pl. in.
nee AML, 6p Qe
Pardaleodes makala Bethune-Baker, Ann. & Mag. Nat. Hist.
ser. 8, vol. ii. p. 480 (1908).
Mr. Bethune-Baker’s type, which he has kindly shown me,
does not differ in any way from Dr. Holland’s excellent figure.
We have several specimens collected by Mr. G. L. Bates at
Bitje, Ja River, Cameroons.
CERATRICHIA IALEMIA, sp.n. (Plate LXVII. fig. 10.)
Male. Allied to C. nothus Fab.* Upper side uniform dark
brown, with 5 or 6 minute white spots beyond the cell on the
fore wing. Under side: fore wing much as in C. nothus; hind
wing white, with 5 small brown discal spots and outer margin
broadly clouded with dark brown enclosing a submarginal series.
of 5 round spots with paler centres. Thorax and abdomen
brown above, white below. Legs white. Palpi yellow with
black hairs.
Female. Upper side as male, but white spots larger and with two.
extra in the discal series towards the inner margin, and a white
spot in the cell of the fore wing. Under side: fore wing
marked as above; hind wing as in male.
Expanse 1,%, inch.
Hab. Bitje, Ja River, Cameroons, 2000 feet (G@. LZ. Bates).
Dry and wet seasons.
This insect has the hind wing below quite differently marked.
from C’. nothus, which is very near to the figure given by
Dr. Holland of Hypoleucis enantia Karsh, P.Z.S. 1896, pl. i.
fig. 17. M. Mabille in Wytsman’s Hesperide still places enantia
in the genus Hypoleucis.
EXPLANATION OF PLATE LXVII.
zl
ae
. Ortholexis melichroptera Karsch, p. 407.
. Abantis ja, sp. n., _p. 408.
3 lucretia, sp. n., p. 409.
. Acleros sparsum, sp. 0., p. 410.
s» kasai, sp. n., p. 410.
Gorgyra afikpo, sp. u., p: 411.
. Parnara noctula, sp. n., p. 411.
ch palocampta, sp. n., p. 411.
i flavifasciola, sp. n., p. 412.
. Ceratrichia ialemia, sp. n., p. 413.
. Celenorhinus nigropunctata B.-Baker, p. 407.
Sj
* Papilio nothus Fab. Mant. Ins. p. 88. n. 799 (1787).
414 ON TICKS CAUSING A RELAPSING FEVER IN MAN. [ Apr. 27,
April 27th, 1909.
Prof. E, A. Mrncuin, M.A., Vice-President,
in the Chair.
The Secretary read the following report on the additions made
to the Society’s Menagerie during the month of March 1909 :—
The number of registered additions to the Society’s Menagerie
during the month of March last was 157. Of these 96 were
acquired by presentation, 21 by purchase, 28 were received on
deposit, and 12 were born in the Gardens.
The number of departures during the same period, by death
and removals, was 135.
Amongst the additions special attention may be directed to :-—
One White-headed Ratel (Mellivora signata), from Sierra
Leone, new to the Collection, deposited on March Ist.
One Eland (Zaurotragus oryx) 5 , from South Africa, presented
by Vice-Admiral Sir Percy Scott, K.C.V.O., on March 9th.
One Eland (Zaurotragus oryx) 9, born in the Menagerie on
March 17th.
One Tasmanian Wolf (Zhylacinus cynocephalus) 2, from
Tasmania, purchased on March 12th.
Two King Birds of Paradise (Cicinnurus regius) 3 2, from
New Guinea, purchased on March 23rd.
One South-African Harrier (Cirews ranivorus), one Booted
Eagle (Visaétus pennatus), and fifty-four Millipedes belonging
to two species (Archispirostreptus pyrocephalus and A. cristulatus),
from Port Elizabeth, presented by A. W. Guthrie, Esq., F.Z.S.,
on March 20th.
Three Hercules Beetles (Dynastes hercules), from Dominica,
new to the Collection, presented by T. Laurance Eve, Esq., on
March 31st.
Mr. H. F. McShane exhibited a series of lantern-slides of
animals living in the Society’s Gardens.
Prof. E. A. Minchin, M.A., V.P.Z.8S., exhibited living speci-
mens of eggs, larve, and adults of the tick Ornithodoros monbata
Murray. This tick lives in the mud floors of human habitations
and encampments in many parts of Africa, and is the means of
transmitting the spirochete which causes African Relapsing
Fever (Spirocheta duttoni). The specimens shown were collected
and sent home by Sir David Bruce, from Uganda, and were
probably infected.
Mr. R. H. Burne, F.Z.S., exhibited a series of specimens
from the Museum of the Royal College of Surgeons, showing
(1) the different mechanisms for the movement of the nictitating
1909.] PHOTOGRAPHS OF BURCHELL’S AND WAHLBERG'S QUAGGAS. 415
membrane, (2) the coarse anatomy of the tapetum lucidum,
(3) the organ for elevating the eyes in the Plaice.
Mr. R. I. Pocock, F.L.S., F.Z.S., the Superintendent of the
Gardens, exhibited the photographs of two Quaggas, more com-
monly called Burchell’s Zebras, which were recently imported
from Zululand by Herr Reiche, to whom Mr. Pocock was indebted
for the photographs. He remarked :—
“These two photographs represent two distinct races of the
Quagga, using that term in its broad and correct sense *, both of
special interest. The first (text-fig. 48) belongs to the race described
by Gray as Hqwus burchelli, which should be called Burchell’s
Text-fig. 48.
Burchell’s Quagga (Equus quagga burchelli).
Quagga (Hqwus quagga burchelli). Very few examples of this race
appear to be in existence, either as living animals or museum spe-
cimens. Ina wild state it is either verging on extinction or wholly
extinct. It was originally recorded from British Bechuanaland,
and there seems to be no reliable evidence of its occurrence
elsewhere. No great importance can be attached to the fact that
the specimen in question is alleged to have come from Zululand,
because, as the photograph shows, the animal wears a head-stal
* Pocock, Ann. Mag. Nat. Hist. (7) x. p. 306, 1902.
416 MR. R. I, POCOCK ON PHOTOGRAPHS OF (Apr. 27,
and has obviously been tamed, and may have been transported to
various places in South Africa. A point to be noticed in the
photograph is the wavy nature of the stripes on the hind-quarters
and the unusually marked indication they show of the tendency
to break up into spots, which is carried to a greater extreme in
the typical Quagga. Shadow-stripes are just traceable up to the
withers, and a few well-defined ones are observable on the neck
close to the mane.
“This animal supplies one more link in the chain of evidence
showing that the true Quaggas and so-called Burchell’s Zebras
are but local races of one and the same species.
“ Similarly the second photograph (text-fig. 49) shows an animal
almost intermediate in pattern between Burchell’s Quagga and
Text-fig. 49.
Wahlberg’s Quagga (Hquus quagga wahlbergi).
Chapman’s Quagga, and supplies a stumbling-block in the way of
those zoologists, if any remain, who hold that a specific distinction
exists between H. quagga burchelli and EF. quagga chapmanni.
The animal depicted is a specimen of the little-known Wahlberg’s
Quagga (L. quagga wahlbergi), which I described some years ago
1909.] BURCHELL'S AND WAHLBERG'S QUAGGAS. 417
from a specimen from Zululand in the British Museum. It is not
exactly like the type, but is near enough to leave little doubt as
to their subspecific identity. It differs from Burchell’s Quagga in
being striped, though sparsely, on the outsides of the legs and on
the hind-quarters up to the root of the tail, and in the extension
of the body-stripes to the median ventral line. The stripes are
all rather thin, and are especially thin and faint on the area below
a line joining the stifle-joint and the root of the tail. On the
lower half of the thigh, on the left side, they form a more or less
reticulated pattern, as has been already recorded in the case of
the example of this race in Mr. Rothschild’s Museum.at Tring *.
On the thigh of the right side the stripes form no such reticu-
lation. Shadow-stripes are well developed and extend as far as
the withers, as in the typical specimen. There are indications of
them also on the neck. Since Zululand is known to have been
the home of Wahlberg’s Quagga, there is no reason to doubt that
the specimen under discussion came from that country.
“Tn connection with these photographs attention may be drawn
to a point connected with the pattern of these and more northern
races of Quaggas, which I believe has a procryptic significance not
previously noticed. This is the sharp division of the body by the
direction of its stripes into an anterior and a posterior half. On
the shoulders and fore part of the body the stripes are vertical ;
but on the hinder half of the body and quarters they abruptly take
an oblique direction backwards. The optical effect this change of
direction in the stripes has upon me is that of two distinct objects
in the line of vision instead of one; each object is in itself in-
complete, and the resemblance of the two combined to the body
of an animal shaped like a uniformly-coloured horse tends to be
destroyed. This obliteration of the horse-like form is still further
effected by the nature of the stripes on the posterior half of the
neck, which are not only broader than those on the withers and
shoulders, but when the neck is carried in its usual position, incline
at a different angle from them. Coming finally to the head, it is
noticeable that there is a further change in the stripes, which
are not only narrow but vary in direction, the direction being
to a great extent different from that of the neck-stripes. As
for the legs, they are either unstriped in Quaggas or striped in
a direction parallel to the long axis of the body. Thus setting
aside the legs, a Quagga of the type above discussed, of which any
one of the subspecies would do as an example, is broken up by the
size and direction of its stripes into four distinct blocks or areas,
namely, the head, the neck, the shoulders and fore part of the
body, the hinder part of the body and the hind-quarters. The
optical disruption of the equine form caused by the continuity of
narrowly striped and broadly striped areas depends upon the fact
that the broadly striped areas appear to be nearer the observer
than the narrowly striped areas. They seem to be two objects
* Pocock, Ann. Mag. Nat. Hist. (6) xx. p. 45, 1897.
418 PHOTOGRAPHS OF BURCHELL’S AND WAHLBERG'S QUAGGAS. [ Apr. 27,
marked in the same way, but one is remote and the other closer
at hand. To detect this the Zebra must be regarded from a
moderate distance and not at so great a distance that the stripes
blend into a uniform grey, as is said to be the case*.
Text-fig. 50.
Sevag *
=
a
x
a
Diagram of one of the northern races of Hquws quagga, to show how variation in
the direction and the width of the stripes contributes to the procryptic effect
of the pattern.
‘Other striped species of the genus Hquus exhibit similar phe-
nomena, but none shows the division down the middle of the side
which is so marked a characteristic of the existing forms of
LH. quagga. In £. zebra and £. foai the change of direction does
not take place until the hind-quarters are reached; but on this
area in /, zebra there is not only a difference in direction of the
stripes, but an extraordinary difference in their size, so that the
greatest possible contrast is secured. Finally, of all the Zebras
HL. grevyiis perhaps the most uniformly striped, but the small
striped head is severed from the small striped shoulder by the
neck, the stripes of which, at least in its posterior half, are usually
of comparatively enormous width. But in the foal of this species
what may be described as an attempt at the optical division of the
body into two halves is made. This is effected not by an altera-
tion in the direction of the stripes, but by an alteration in their
* There is, in my opinion, very little doubt that the sharp contrast im colour
between the black anterior and white posterior parts of the body of the Malayan
Tapir (Lapirus indicus) has a procryptic significance similar to that achieved by
the alteration in the direction of the stripes seen in existing forms of Quaggas.
1909.] THE SPECIES OF THE LEPIDOPTEROUS GENUS LycHNorsis. 419
colour, the stripes on the cheek, neck, and shoulders bemg black
and sharply contrasted with those on the posterior part of the
body and quarters, which are rusty brown. This at least was the
case in the only foal of this species that I have seen, namely, one
born in the Gardens. Another interesting feature about this little
animal was the presence of an erect, thick mane, mostly rusty
brown in colour, which was continuous with the neck-mane on the
withers and extended backwards down the spinal stripe, over the
tail to the tail-tuft. And if, as appears highly probable, Prof.
Ewart is correct in thinking Grévy’s Zebra the most primitive
type in the matter of pattern, it may be that this transitory spinal
mane in the foal of that species is evidence that the ancestor of
all Zebras, and inferentially of all Equide, since 2. grevyi is, in
my opinion, proved by his braying voice to be nearly akin to the
African Wild Ass, possessed a mane running from the crown of
the head to the tip of the tail.”
The following papers were read :—
1. A Review of the Species of the Lepidopterous Genus
Lycenopsis Feld. (Cyaniris auct. nec Dalm.) on exam-
ination of the Male Ancillary Appendages. By T.
ALGERNON CHAPMAN, M.D., F.Z.S.
[Received March 17, 1909.}
(Text-figures 51-122.)
This genus has been generally known as Cyaniris Dalman,
since Scudder so defined it in 1872. Prout, however, calls
attention to the fact that Dalman himself gave semiargus
(argianus) as the type, a circumstance that Scudder in some way
overlooked.
Felder, in 1865, gave Lycunopsis as a generic name to haraldus
(ananga), and there is now a fair consensus of opinion, which the
observations reported in this communication fully support, that
haraidus is a species of Cyaniris (Scudder). Mr. Prout tells me
that notwithstanding Felder gave the name Lycwnopsis to express
his opinion that haraldus was not congeneric with anything else,
the rules at present accepted make Lycenopsis the generic name
of Cyaniris (Scudder). Mr. Tutt, in 1906, unaware of the claims
of Lycenopsis to this position, provided the name Celastrina
with argiolus as type. Should some one find reason to divide
Lycenopsis into several genera, Celastrina would become the
name of that containing argiolus, I do not myself see, at present,
any reason for such subdivision of Lycenopsis, and could not
frame divisions, without using as generic characters features that
seem to me to have only specific value. It remains to remark
that, as Cyaniris has had a start of only 30 years, it hardly
claims place under the new agitation for advocating usage v.
420 DR. T. A. CHAPMAN ON THE SPECIES [Apr. 27,
priority. Such new practice, moreover, would only be binding
after each case was determined by a Committee, appointed by the
Zoological Societies of the world. Short of this it would only
prolong the confusion due to a struggle ultimately to be decided
in favour of priority.
I was led to examine this group by Mr. Tutt desiring me to
investigate Z. argiolus. I found this could not be done “without
dealing with a number of other, chiefly Indian, species, and these
again required, or at least suggested, a still wider view, so that I
finally took up the whole genus, so far as I could obtain access
to material. No doubt, my view is still narrower than it ought
to be, because I have not gone fully into the most nearly allied
genera. I have, in fact, made observations of some of these,
but it was obvious that I must draw the line somewhere and take
the risk of such errors as ignorance of what was beyond it might
lead me into.
Another serious limitation was found in the fact that specimens
of some species could not be obtained for examination, and this
unfortunately was apt to affect most especially type and other
more valuable specimens. Still, I have examined a great deal
of material, but there are six species of which I know little, of one
of these only the 2 is known, and several of the others seem to
be probably synonyms.
I have examined a. good many species in the British Museum
Collection at South Kensington, and am also under great obligation
for the privilege of examining many specimens from the Rothschild
Collection at Tring. Mr. H. H. Druce has afforded me assistance
with specimens and aid in determining species. Mr. Bethune-
Baker has kindly allowed me to examine specimens including
several types from his collection, JI have also had specimens
from the Godman Collection, from that of the Oxford University
Museum, and elsewhere. The late Col: Bingham also most
kindly assisted me with specimens and afforded me the valuable
aid of the researches he had made in writing the volumes on the
‘“‘ Butterflies” in the ‘ Fauna of British India.’
I have. found considerable difticulty in making sure that the
specimens I examined were correctly named, and in the absence
of access to types of well-known species, the authority of the
works of Moore, de Nicéville, and Bingham, for the Indian species,
was generally found to be adequate.
My observations of the genus lead me to the conclusion that
the upper surface is often more misleading than otherwise as to
what species some particular specimen belongs to. As to the
under surface, I believe the pattern of the spotting is to be
thoroughly depended on for specific characters. But then it
unfortunately happens that the pattern is not unfrequently quite
obscured by seasonal, geographical, and other variations, the spots
being much enlar, zed, or some of them reduced almost or quite to
obsolescence. They may be very dark or very pale, and so on.
It has happened to me several times that I saw no identity
1909. ] OF THE LEPIDOPTEROUS GENUS LYCHNOPSIS, 421
between the spots of two specimens of the same species until an
examination of the appendages led me to believe they might be
the same species, and with this indication an identity of the
arrangement of the spots in both could easily be accepted.
Considering these difficulties, it is not to be wondered at that
some confusion has, hitherto, existed as to certain species. The
wonder is that the amount of accuracy, that actually obtains, has
been achieved. I would, however, sincerely beg no one to name
any more “species” until he has fairly located them as new or
otherwise, by examining the appendages, which give an uncertain
answer in very few cases,
What appealed to me as perhaps the most interesting circum-
stance I have unearthed, is the remarkable (apparently mimetic)
variation that occursin LZ. argiolus. Possibly the hope of meeting
with similar cases urged me to continue the researches. Though
other species present considerable variations of the same character,
their number and range is far short of that of argiolus. It
remains, however, very possible that many more similar cases
are still to be discovered when larger series of specimens from
the different Malayan islands are available for full discussion.
I have not found any account of the ancillary appendages of
this group, but I am not a successful searcher of literature; still,
I believe no such account exists. Some observations have, how-
ever, been made, as I find that Doherty (J. A. S. B. 1886, p. 134)
refers to the value of the “ Prehensores” for distinguishing the
species of the group. I cannot discover, however, that he
afterwards published any details.
Though I have examined the ancillary appendages of various
Lepidoptera of many different families, I am much impressed
with the fact that my knowledge is really very superficial and
fragmentary. The result is, that in defining the generic characters
of these organs in Lycenopsis, | do not experience very much
confidence.
Throughout the Lycznide it is hardly too sweeping a state-
ment, so far as my observations go, to define the dorsal armature
as consisting of two lateral halves with more or less of a hiatus
between them, each lateral half carrying near its base a paramere
or appendage, articulated to 1t so as to be freely movable and more
or less curved, bent, or twisted. In a few genera the two lateral
halves coalesce in the middle line. In Lycenopsis we have a
remarkable specialization of these dorsal processes, a specialization
by way of simplification or loss of parts. The movable paramere
or hook is quite wanting.
This specialization 1s so remarkable as at least to justify the
inclusion in the Lycenopsids of all genera that show this special-
ization, of which at present I have only recognized Castalius,
Megisba, and Neopithecops. Of those genera apparently near
that possess these hooks, such as Pithecops, Niphanda, &c.,
the only one that I incline to regard as Celastrinid is Zizera,
422 DR. T, A. CHAPMAN ON THE SPECIES [Apr. 27,
[type maha, not alsus (minimus); alsus is an Everid and was
stated to be the type of Zizera by Moore under some misappre-
hension. It does not possess the characters by which he defines
the genus, whilst maha (and other Indian species) does].
Two species placed in Lyeenopsis, viz. vardhana and musine
(Jugra), possess these hooks, and probably require each a separate
genus, but I shall satisfy myself for the present by placing them
in the genus Votarthrinus, instituted for an apparent Lycenopsis,
possessing this character. catreus, rather an aberrant form,
would be just within Lycenopsis. Before inventing genera for
these, it would be well to know something of their life-histories.
This tolerably uniform structure of the dorsal armature
throughout the true Lycnopsids reinforces the remark I made
a number of years ago, in discussing the morphology of these
structures in the genus Hrebia. I then observed that the dorsal
appendages were fairly constant throughout a genus, whilst the
clasps varied very much, and yet very similar variations in the
clasps could be found in allied genera clearly marked off by the
dorsal structures.
These lateral hooks of the dorsal armature must be, from
their constancy throughout the Lycenids, of some essential use,
which we must suppose was not imperative in the Lyceenopsids.
Nevertheless, in Lycenopsis we find that some species apparently
regretted the loss, but under the law that lost parts cannot be
regained, could only meet their desires by developing a more or
less hard and sharp point as part of the lateral dorsal process,
most highly developed in LZ. limbata (19).
In the few notes I may make on theappendages of each species,
I certainly do not attempt any full description, trusting, so far as
that goes, to the photographic reproductions. In such remarks
as I may hazard, my objects are rather to call attention to points
that mark their agreements or differences with other species, as
hints to their affinities, and to points not clearly brought out by
the photographs.
In preparing the specimens, I found that to secure their being
satisfactorily photographed and easily compared under similar
conditions, it was necessary to make them quite flat and all
in the same manner: this is true of preparations of ancillary
appendages of most groups, but the question varies in each group
as to how such flatness may best be attained. In some groups it
is well nigh impossible. In. the Lycenopses I find it fairly
satisfactory to divide the basal chitinous ring in the medio-dorsal
line and spread it out, so that the extreme ends of the preparation
are the separated dorsal portions of the ring. The photographs
appended are nearly all from such specimens. They are mostly
magnified 45 diameters, more or less reduced on reproduction,
but there are a few exceptions. The photographs are by Mr. F.
Noad Clark, whose technique leaves nothing to be desired, except
specimens mounted in a more finished and less amateur manner
to operate on.
1909. ] OF THE LEPIDOPTEROUS GENUS LYCNOPSIS. 423
In one or two instances specimens are mounted so as to show
the dorsal armature in more natural relations, as one of dilecta
(23), of selma (29 a).
It is a matter of congratulation from my point of view, that
the three species that are especially widely distributed and of
most protean aspect, viz., argiolus, puspa, and limbata, have
ancillary appendages that are absolutely distinctive and widely
marked off in each case from those of any other species and are
also given to very little variation, geographical or other. It is
different with the dilecta group of species which do not differ
widely from each other, and amongst which I think it possible
there are still one or two undetected species, but my information
is too imperfect to enable me to deal adequately with this
section.
I think the latest complete list of Lycenopsids is that by Dr. A.
G. Butler in the ‘ Annals and Magazine of Natural History’ for
1900, ser. 7, vol. v. p. 441. It is curious that though de Nicéville
(1890) characterises the genus correctly, Dr. Butler should define
it by one character only, and that one, one that it does not possess
—the costal vein of the primaries is not united by a cross-vein to
the first subcostal branch. This cross-vein is rare in Lycenines,
and it is absent in all the genera having Lycenopsid affinities,
It is present in the Everids and some few other genera,
Dr. Butler enumerates 57 Cyanirids and 3 doubtful species. To
these 60 species there are in addition several further species de-
scribed since and one or two he had overlooked, making 66 in all.
Of these two are Everids (Lothrinia), three belong to Notarthrinus,
and perhaps are hardly Celastrinid, one (pryeri) requires a new
genus (Artopoetes), leaving 60 to Lycenopsis. Of these 60 forms
I have not examined 7 or 8, but believe several of them would be
sunk under other names were that done. The remaining 52
appear to belong to 30 distinct species, 22 being usually varietal
forms rather than synonyms, but in either case included in the
30 “ good ” species.
I have added photographs of examples of the three genera that
appear to me to be undoubted Lycenopsids—namely, 45, Megisba
malaya, 46, Neopithecops zalmora, and 47, Castaliusethion; it will
be observed that in each of these the dorsal armature is without
an accessory hook.
I may begin with those species that I separate from Lycenopsis
under the genus Wotarthrinus, of which the definition is appa-
rently Lycenopsis, but with hooks to the dorsal processes of the
appendages. Of these there are three species, vardhana, binghami,
and musina, sufficiently different from each other to be doubtfully
accommodated together in the same genus. In the boldness of
the teeth of the clasps, musina seems related to the group of
Lycenopsis containing nedda, shelfordi, corythus, &e. Tf this be
so, this group would probably be the earliest or lowest species of
the genus.
424 _ DR. T. A, CHAPMAN ON THE SPECIES [ Apr. 27,
NOTARTHRINUS,
1. vardhana Moore, P. Z. 8. 1874, p. 572, pl. Ixvi. fig. 5.
The dorsal armature possesses the accessory articulated hooks,
so that it is not a true Lycewnopsis. I place it provisionally in
Notarthrinus.
Text-fig. 51.*
vardhana. X33.
2. binghami Chpm. P. Z. 8. 1908, p. 676, pl. xxxviu. fig. 2.
Very like Bothrinia chennellii and Lycenopsis limbuata var.
jynteana, and until recently not distinguished from them. The
appendages are very different.
(Appendages illustrated J. ¢.) :
3. musina de Nicév. J. Bomb. N. H. 8. ix. p. 275, pl. O. fig. 19.
& Specimens of this species in the British Museum collection are
also labelled corythus—leading me at first into some errors,—
specimens of corythus, however, exist there unnamed.
} This species is on the margin of, or rather outside, the genus
Lycenopsis. It possesses articulated hooks to the dorsal pro-
cesses, and the terminal armature is differently arranged to that
of any other species. In its large and bold spines it resembles
corythus, cinctata, and transpecta, and some others, most of which
* [Text-figures 51-122 represent in each case the male ancillary appendages of the
species indicated in the legend. |
1909. | OF THE LEPIDOPTEROUS GENUS LYCNOPSIS. 425
Text-fig. 52.
musina (labelled corythus, B.M.). X 38.
Text-fig. 53.
musina (B.M.). X33.
Proc. Zoou, Soc.—1909, No. XX VIII. 28
426 DR. T, A. CHAPMAN ON THE SPECIES [Apr. 27,
Text-fig. 54.
musina (lambi, Oxford Coll.). X 41.
Text-fig. 55.
musina (lugra, Druce Coll.). > 41.
1909.3 OF THE LEPIDOPTEROUS GENUS LYCHZNOPSIS. 427
have also a soft (not jointed) small appendage on the ventral side
of the dorsal process, that probably represents the obsolete hook.
If this be so, muwsina would be an ancestral form, hardly yet a
Lycenopsis, and the others referred to would be the earliest true
_Lyceenopsids.
3 Ge Unga aly lel, 1Dienee. 12. Ao tee isa), jos DUH poeany ines, G,
has identical appendages and differs from musina most
in the upper surface-colouring. It is to be regarded as a
geographical race of musinda.
Text-fig. 56.
musina (lugra, Tring Coll.). X 41.
Lyc2NOPSsIs.
Tn classifying the appendages, there are no grounds apparently
for supposing one can easily at the same time classify the species.
One is inclined to begin with puspa as almost the only species
that can be said to have a hard (spinous) and soft (hair-clothed)
process to the clasps, because this seems to be not only a primitive
Lycnine but even a primitive lepidopterous character, following
with argiolus as possessing vestiges of the soft process. In other
species the soft process is merely constructively present. After
these would come such species as oreas, phillippina, &e., in which
the spinous process bends down, and in corythus, nedda, &e.
displays the teeth as marked spines.
I8*
a
428 DR. T. A. CHAPMAN ON THE SPECIES (Apr. 27,
In another direction the hard or spinous process dwindles to
small dimensions, retaining many teeth, as in dilecta, lanka, &e. ;
culminating in species in which it is very small, as in placidula,
melena, &c.; and finally wanting, as in plauwta. ‘There are other
species that seem each to be a law to itself, such as albocwrulea
and transpecta.
A group might be made of those that have, not a hook, but a
spine to the dorsal process; it would not, in regard to the clasps,
be a natural group. It appears to include Limbata, marginata,
phillippina, nedda, acesina, camene, argiolus.
T propose to place first those with several large teeth to the
spinous process.
4. corythus de Nicév. J. B. N. H.S8. ix. p. 2738, pl. O. figs. 16,
17.—Sumatra.
The appendages belong to the group with long teeth in the
spinous process of the clasps—nedda, acesina, shelfordi, musina.
The dorsal processes have a definite spine and a button-like
ventral accessory.
Text-fig. 57.
corythus (Sumatra, Druce Coll.). XX 32.
In the British Museum specimens of musing are placed under
this name, which led in my earlier examinations to my believing
corythus and musina to be the same species (Proc. Zool. Soe.
1908, p. 677). They are abundantly different in all respects.
1909. ] OF THE LEPIDOPTEROUS GENUS LYCNOPSIS. 429
5. shelfordi de Nicév. J. B. N. H.S. xiv. p. 245, pl. FF. fig. 7.—
Borneo,
The dorsal process is without a spine, the ventral accessory is
flat and rounded but well chitinised; the clasp is short and the
end broad, with very strong and jong spines, the secondary ones
being strongly developed. One might describe it as a strong-
spined form like nedda trying to shorten itself down to the form
of placidula.
Text-fig. 58.
shelfordi (Druce Coll.). X 40.
6. nedda Grose-Smith, Nov. Zool. 1. p. 572.
6 a. cinctata Grose-Smith, Noy. Zool. 11. p. 506.
66. phuste H. H. Druce.
nedda and cinctata appear to agree in imaginal characters ex-
cept as regards the width of the dark border. Whether they are
geographical or seasonal varieties I do not know. I have, under
phuste, given my reasons for considering this form probably an
aberration of nedda.
A specimen labelled aga in the Godman Collection (now B.M.)
and one labelled ananga in the Brit. Mus. Collection belong to
this species.
The dorsal processes have a short sharp ventral spine; the
clasps are short and broad and have the outer face of the spinous
process facing distal, with very large bold teeth, 4 or 5, culmi-
nating in a very strong epeninn | one; the noes beneath is
segues
430 DR. T, A. CHAPMAN ON THE SPECIES [ Apr. 27,
Text-fig. 59.
nedda (cinctata). X 43.
1909. | OF THE LEPIDOPTEROUS GENUS LYCENOPSIS. 431
Text-fig. 61.
nedda (aga, Godman Coll.). x 44.
Text-fig. 62.
nedda (labelled ananga, B.M.). X 44.
432 DR. T. A. CHAPMAN ON THE SPECIES [Apr. 27,
7 (66). phuste H. H. Druce, P. Z.S. 1895, p. 573, pl. xxxiv.
me, Y/
I have seen the type specimen, which is I believe unique. Not
having examined the appendages I can make no positive assertion,
put I see no difference between phuste and cinctata except the
absence on the under side of all spots except the marginal ones.
This isa form of aberration one expects to meet with occasionally
in most Lycenines. Ordinary cinctata occurs in Dili, whence this
specimen comes.
8. cara de Nicév. J. B. N. H.S. xii. p. 143, pl. Z. figs. 19 & 20.
T have not had an opportunity of examining this species. It
seems related to acesina.
9. acesina Bethune-Baker, Ann. Mag. N. H. ser.7, xvi. p. 103;
IP. ZA, Sy, UDO, Jo, WIG, jails switinis saise, 10),
The appendages here are very distinct. It is not far from
nedda in general structure, but it is even bolder and more spinous.
The ventral spine of the dorsal process is very strong and sharp.
The spinous process of the clasp is separated off as a head with a
distinct neck, the base carrying the inferior spines being much
enlarged (as compared, say, with nedda), and pr ojecting some
distance beyond the large bold terminal spine. In musina this
development is carried further.
I have not seen Z. cara, but the figures suggest it is closely
allied to acesina, possibly the Same species.
10. transpecta Moore, P. Z. 8. 1879, p. 139.
The appendages differ much from those of any other species of
Lycenopsis. The dorsal process differs from that typical of the
genus in having a soft ventral process. The clasps have a wide
unarmed extremity, but mar ked teeth exist along the ventral
margin. corythus is the nearest form ; musina thas some similar
features (see musinda).
The soft ventral process of the dorsal armature is probably a
vestigial remnant of the base of the typical hook, and may mark
the species possessing it (corythus, catreus, wc.) as earlier forms.
11. phillippina Semper, Reise Philipp. um. v. p. 168.
A specimen in the Godman Collection (now in Brit. Mus.) is
almost certainly correctly named. The appendages are very
distinct from those of any other species. The dorsal process has
a sharp ventral point. The clasps are short, very broad and stout,
and end by continuous tapering, but with a sharp curve, in a
strong spinous process with teeth along the distal edge and end-
ing in a large robust point. It is most “nearly allied 40 oreas.
1909. ] OF THE LEPIDOPTEROUS GENUS LYCHNOPSIS. 433
acesina. X 42.
Text-fig. 64.
Mage ak
PA
Mb ‘i ait Ey x a
434 DR. T. A. CHAPMAN ON THE SPECIES [Apr. 27,
Text-fig. 65.
phillippina (Godman Coll.). X 42.
12. oreas Leech, Butt. Chin. p. 321, pl. xxxi. figs. 12-15.
This is a well-marked species. The clasp somewhat resembles
that of phillippina, but is perhaps nearer to puspa or nedda in so
far as that the inner angle (soft process) instead of sloping away
is fairly produced; the terminal spine is large and sharp, those
below it are mere notches—smaller and smaller as they recede, to
the number of six or eight. The dorsal process has a ventral
spine.
There is a specimen in the British Museum labelled singalensis,
Khasia Hills. This may be one of the specimens mentioned (from
the Nilgiri Hills) by de Nicéville, Butt. Ind. iii. p. 108, and adds
oreas to the Indian fauna unless there be some error in labelling,
which I see no reason to suspect.
13. haraidus Fabr.
The clasps are broad and short, like phillippina, oreas and ripte,
but are larger, and the spine is larger than in ripte, having large
and bold teeth, and as long as the width of the clasp. The dorsal
process has a small accessory on its ventral side.
14. ripte H. H. Druce, P. Z. 8. 1895, p. 574, pl. xxxi. fig. 11.
In form of clasp this species is very near to placidula, excepting
that the spinous process is well developed, not only extending
1909. ]
OF THE LEPIDOPTEROUS GENUS LYC/NOPSIS.
Text-fig. 66.
oreas (China), X 42.
Text-fig. 67.
oreas (labelled singalensis, Khasia Hills, B.M.).
x 32.
436 DR. T. A, CHAPMAN ON THE SPECIES [ Apr. 27,
Text-fig. 68.
haraldus. X 44.
Text-fig. 69.
ripte (Godman Coll.). X 41.
1909. | OF THE LEPIDOPTEROUS GENUS LYCANOPSIS. 437
two-thirds of the width of the clasp across its end, but having a
deep notch below it and having five large marginal teeth, gradu-
ated larger to the terminal one, which is strong and bold,
haraldus is very near it, but is larger and more robust.
There is no dorsal accessory process.
Text-fig. 70.
ripte (Oxford Coll.). X 37.
15. puspa Horst. Cat. Lep. E. I. Co. p. 67. n. 3 (1828).
This seems to be the most abundant and most widely dis-
tributed species after argiolus. Like argiolus it has received
in several varieties distinct names, in the belief that they were
distinct species. Its distribution appears to be over practically
the whole Indo-Malayan region, and we know that in many of the
islands the species is found, often with a very definite varietal
facies, and it is probable that nearly every island in the region
possesses a race of puspa.
puspa may always be recognised by the spots of the upper
wing (beneath) having those between veins 2 and 4 and 4 and 5
oblique, with the intermediate ones fairly straight and nearly in
the line joining the centres of the oblique ones, and that between
4 and 5 on the hind wing being similarly longitudinal rather
than transverse. :
The appendages are nearer to those of argiolus than those of
any other species. The very similar spiculate spine, instead
of having four or five teeth, has a very large number of very
minute ones. The body of the clasp is also very much longer
proportionally.
438
DR. T. A. CHAPMAN ON THE SPECIES [Apr. 27,
The following recognised (?) species are apparently varieties
of puspa :—
timorensis Butl. A. M. N. H. v. 1900, p. 443. Timor.
dammce Heron, A. M. N. H. xiv. 1894, p. 104. Damma I.
splendens Butl. A. M. N. H. v. 1900, p. 444. Perak.
kuhni Rober, Isis, 1. p. 60. Ke.
imperatric Butl. A. M. N. H. v. 1900, p. 444. Siam.
cajaya Felder, Reise Novara, Lep. 11. p. 278. Luzon.
cyanescens de Nicéville, Butt. Ind. iu. p. 103. Nicobar.
puspinus Kheil, Rhop. Ins. Nias, p. 30. Nias.
lambi Distant, A. M. N. H. ser. 5, xix. p. 266.
Of these, eyanescens has a coloration that makes one doubtful
of its identity with puspa until an examination of the appendages
shows them to be truly the same species.
Amongst other forms (and localities), some of which may have
been named though I have not discovered it, and for each of
whi
ch I would suggest, should a varietal name be desired, a
varietal name indicating the habitat, may be mentioned :—
1
y
Or
“I
. A large form (34 mm.) with large white patches on all four
wings, and very white under side on which the usual spots
stand out very black and crisp. From Sumbawa. Tring
Coll.
2. A form almost identical with cyanescens, but with the special
tint of that form less pronounced ; under side with a grey
tone. Rawau. Tring Coll.
. A moderate-sized form (30 mm.) with white patches indi-
eated on all wings ; under side a little clouded. From Celebes.
Tring Coll.
- Moderate size (33 mm.), broad border, no white patches
or discal streak ; under side grey-clouded. Battak, N.E.
Sumatra. Tring Coll.
. (84 mm.) Narrow border, white patches hardly indicated ;
under side clouded grey. Bungurun, Natuna Island. Tring
Coll.
. A very narrow bordered form (35 mm.), white patches
hardly indicated, slightly clouded beneath. Palawan.
Tring Coll.
. A fairly ordinary form with white patches. From Sawela,
Lombok. Tring Coll.
. A very dark blue form without white patches. Sula Resi.
Tring Coll.
Miskin mentions that it occurs in Australia, but without
quoting definite data. G. A. Waterhouse is doubtful whether it
does so.
1909. ] OF THE LEPIDOPTEROUS GENUS LYCHINOPSIS. 439
Merx=ico(ale
puspa. X 44.
Text-fic. 72.
puspa, end of clasp. X 162.
440 DR. T. A. CHAPMAN ON THE SPECIES [ Apr. 27,
Text-fig. 73.
puspa (imperatrixz ?, Luzon, Tre Coll.). xX 46.
Text-fig. 74.
puspa (cajaya). X 32.
1909. | OF THE LEPIDOPTEROUS GENUS LYCNOPSIS. 44]
Text-fig. 75.
puspa (lambi, Godman Coll.). X 43.
Text-fig. 76.
puspa (duponchellii, Godman Coll.). X 45.
Proc. Zoo. Soc.—1909, No. X XIX. 29
449 _. DR. T. A, CHAPMAN ON THE SPECIES [ Apr. 27,
Text-fig. 77.
puspa (labelled damme, Shevoray, Tring Coll.). x 45.
Text-fig. 78.
puspa (Sumbawa, Tring Coll.). x 44.
1909. j OF THE LEPIDOPTEROUS GENUS LYCENOPSIS. 4435
16. argiolus Linn.
A good deal of what I have done in investigating this species
will be found in Tutt’s account of it (Brit. Lep. vol. ix. p. 387
et seq.), and in Proc. Ent. Soc. Lond. 1908, p. lxxxi.
The number of named varieties, races, and aberrations is very
great. I have examined a very large number of specimens, and
the ancillary appendages are all identical and quite unlike those ot
any other species of Lycewnopsis, puspa being perhaps the nearest.
The amount of variation in the appendages is apparently confined
to the greater or less prominence of the four or five teeth on the
spine of the clasp. They are never entirely obsolete, and are
perhaps as pronounced in British specimens as in those of any
other race. The soft process of the clasp varies a little in its
prominence, though this may depend sometimes on slight varia-
tions of procedure in mounting the specimens.
Text-fig. 79.
argiolus (English). X 37.
I will not deal with varietal names, beyond one or two in
addition to those that I find in Dr. Butler’s list.
Dr. Butler confines the name argiolus to the European form,
to the American form he gives the name ladon Cramer ; Cramer’s
figure looks like a Lycenopsis, and like an argiolus. The assigned
locality is obviously erroneous, but whether the specimen actually
came from Europe, Asia, or America, is quite a matter of doubt.
pseudargiolus seems therefore the name for the American form.
Zor
444 DR. T. A. CHAPMAN ON THE SPECIES [Apr. 27,
There seems to be no other American Lycenopsis, but the species
is found over the whole of North America, from Centra] America
to Canada, with a profusion of named and unnamed races and
aberrations.
The Asiatic forms mentioned by Dr. Butler are :—
levettii Butler. Corea.
huegelti Moore. Himalayas.
ladonides de YOrza. Japan.
celestina Kollar. Western India.
victoria Swinhoe. Shillong.
The last species astonished me a good deal when I first found it
to be a race of argiolus, but its markings, &c. are found to conform
to those of argiolus.
One of the most interesting features of the Indian Lycnopsids
is the parallel variation of certain species, and probably of others
the variability of which has not so directly come under my notice.
The first point to attract my attention was the similar forms
found in Z. argiolus and L. limbata.
We have ZL. argiolus vars. huegelit and cclestina paralleled by
L. limbata and its var. placida. This would not strike one as
remarkable were it not for the further close likeness between
argiolus var. sikkima, and limbata var. jynteana.
These two forms were distinguished by Moore and separately
dlescribed, but he afterwards was obviously unable to name them
with certainty, and there is a specimen of sikkima in the Brit.
Mus. Collection named by him jynteana.
De Nicéville, Butt. of India, vol. ni. p. 105, treated the dis-
tinction between these two species with more than scepticism, and
Bingham sinks sikkima asa synonym of jynteana. Both forms
seem to vary in the breadth of the dark border and in the presence
or absence of a discal line, but sikkima inclines to the broadest
border and is less rarely without the line. Apart from the
ancillary appendages, which show sikkima to be aform of argiolus
and jynteana to be a race of limbata, the structures being
exceedingly different, the two forms may be separated by the
line of spots under the fore wing, the third spot (second of those
in a row, the first being moved inwards) being slightly oblique in
jynteana (as it is in the co-specific forms limbata and placida),
whilst it is like the others in sikkima (as it is in argiolus, though
there are some races in which this varies).
There are two further forms of argiolus that I hardly like to
give separate names to, because they are. really very close to
sikkima, if not identical: one of these flies with Z. puspa. Of
this I have four rather poor specimens received in papers from
Col. Bingham, along with a number of puspa collected by
Col. E. R. Johnson in Assam: submitted to Col. Bingham, he
thought they might be jynteana, puspa, alboceerulea, or chennellit.
They were, as we have seen, jynteana in almost the same sense
that sikkima is jynteana. It is really very close to sikkima, but
1909. ] OF THE LEPIDOPTEROUS GENUS LYCHNOPSIS. 445
Text-fig. 80.
argiolus (pseudargiolus). X 42.
Text-fig. 81.
argiolus (labelled limbata, Godman Coll.). X 40.
446 DR. T. A, CHAPMAN ON THE SPECIES [Apr. 27,
has a rather wider border, and the discal streak has a pale patch
in the blue like puspa. If it must have a name, it might be
argiolus var. puspargiolus.
The other form equally close to sikkima flies with Bothrinia
chennellii, so far as I can judge, and has been sent to me as that
species. It has a still broader border, and would certainly not
suggest itself as a form of argiolus until the ancillary appendages
had been examined. This form might be called bothrinoides.
The first spot beneath the upper wing being thrown inwards (as
in argiolus) easily distinguishes it from chennellir.
There is a further variety of argiolus that I have called albo-
ceruleoides; it is rather further from stkkima than the two
forms ( puspars giolus and bothrinoides) just noticed, but nearest to
puspargiolus in having a whitish patch in the blue of both Wings ;
the black border is, Tarorere, less extensive. There is a specimen
(or more) of this in the B.M. Collection labelled albo-ccer uleus, but
it really is nearer to marginata; and I have specimens of unknown
origin of marginata and albocewruleoides, the pinning and setting
of which suggest that they were taken together. One can hardly
avoid believing that there are probably races of argiolus not yet
observed that approach other species of Lycwnopsis in appearance.
There is a further form, in which the discal line is evident and
the black border narrower, but with distinct traces of lunules on
the margin of the hind wing and very pale in the blue colour.
There are specimens in the British Museum from Burmah, and I
have seen others labelled gynteana, without locality ; but for its
larger size, this is not very unlike victoria, especially in the fading
of the blue into a greyish tone, in which it has not gone far, but
still has moved in that direction.
There is still Lycenopsis victoria Sev., which I found to my
astonishment to be also a race of argiolus. The conclusion can
hardly be resisted that it also has adopted a resemblance to some
other Lycenid, but I am not prepared to suggest what this
may be. :
There are sundry other Lycenopsids that suggest that they
have local races to accord with other species, but I know too little
of these to be able to say more than that such conditions seem
probable. In most of these cases, however, the explanation
of simple geographical variation is at least quite as probable, as in
the case of LZ. eyanescens, the very different looking form that
puspa assumes in the Nicobars.
17. albidisca Moore, P. Z.S. 1883, p. 524, pl. xlviii. fig. 7.
The appendages show this species to be related to marginata as
closely as the general appearance of the imagines might lead one
to suspect.
The spines of the dorsal processes are less pronounced, the
accessory processes on their ventral side are at least equally
distinct; the shoulder of the spinous process of the clasp, so usually
1909. | OF THE LEPIDOPTEROUS GENUS LYCENOPSIS. 447
rounded, is here, as in marginata, sharply angular; the process
itself is smaller, free from the body of the clasp for only a short
distance. The body of the clasp is much more slender than in
margimata.
Text-fig. 82.
albidisca. X 30.
18. marginata de Nicéville, J. A. 8. B. iii. p. 70, pl. 1. fig. 9.
This species seems to be quite distinct and definite in the
ancillary appendages. It belongs to the group in which the
dorsal processes possess a spinous process. The clasps are narrow,
the spinous extremity is small; the shoulder, usually rounded, is
here sharply angular, almost a right angle, so that the, properly
outer, margin is transverse to the axis of the clasp.
A specimen in the Tring Coll. is labelled ‘“‘ Penang, Curtis.”
19. limbata Moore, P.Z.S. 1879, p. 139.
19 a. placida de Nicéville.
19 b. jynteana de Nicéville.
19 ¢. lyce Grose-Smith.
19d. lyseas Grose-Smith.,
(strophis H. H. Druce ®).
Having examined the appendages of many specimens of each, I
448 DR, T. A. CHAPMAN ON THE SPECIES [Apr. 27,
Text-fig. 83.
marginata. X 4).
Text-fig. 84.
marginata (Penang, Tring Coll.). X 42.
1909. ] OF THE LEPIDOPTEROUS GENUS LYCZNOPSIS. 449
Text-fig. 85.
limbata. X 42.
Text-fig. 86.
limbata (labelled Jambi). X 42.
450 DR. 1. A. CHAPMAN ON THE SPECIES [Apr. 27,
am certain that limbata, placida, and jynteana all belong to one
species.
Of lyce I have not examined the type specimen, but another
one from the same locality that agrees with it in every respect.
Of lyseas I have compared the type specimen with forms of
limbata from several Malayan islands, and have no doubt of its
identity.
strophis I have not seen: I throw out the suggestion that it
may be a form of limbata.
The appendages of this species are so characteristic and different
from anything else, that the identity of the specimens examined
is unmistakable. The dorsal process has a very long spine, and
the spines of the clasps incurving and ending in several teeth
(instead of one terminal one) are matched by no other species.
The recognised distribution of limbata, placida, and jynteana
does not extend beyond continental India further than Ceylon and
Penang.
Besides the type specimens of lyce and lyseas, the Tring Coll.
possesses specimens from 8. Flores, 8. Celebes, Upper Aroa River,
N. G., Lombok, North Formosa, N.E. Sumatra; the latter
specimen is a curious dwarf only 23 mm. in expanse.
Most of these specimens are not far from limbata in appearance,
but those from 8. Celebes and Upper Aroa River have wider
borders, without approaching or at least reaching the jynteana
form.
Some remarks on this species in connection with the probably
mimetic significance of the jynteana form, will be found in Tutt’s
Brit. Lep. vol. ix. p. 408, and in Proce. Ent. Soc. 1908, p. lxxxi.
Specimens from W.C. Sumatra are in the Druce Coll. They
probably deserve a varietal name.
limbata and placida from the Moore Collection are both one
species; but in the placida series the spots beneath the fore
wing are distinctly en échelon, whilst those of the limbata are
much more in a continuous line.
19. beretava Ribbe, Deutsch. ent. Zeitsch., Lep. 1899 (Iris)
p. 243, pl. iv. fig. 14.
So far as it 18 possible to judge from a figure, I think this is
probably a form of limbata.
20. strophis H. H. Druce, P. Z. 8. 1895, pl. xxxii. fig. 4, p. 573.
This species has not been examined. I entertain a strong
suspicion from the figure (J. c.) that like lyce and lyseas it may be
a variety of limbata. Mr. Druce’s suggestion that it is a
seasonal form of dilectissima is also possible. The marginal line
enclosing the marginal spots, always fairly well marked in all
the forms of limbata I have seen, is quite wanting. The arrange-
(=)
ment of the spots is otherwise very nearly identical in limbata
1909. ] OF THE LEPIDOPTEROUS GENUS LYCEHNOPSIS. 45]
and dilectissima. This marginal line varies in argiolus, tenella, and
other species from absence to considerable prominence.
21. coalita de Nicéville, J. B. N. H. S. vi. p. 363, pl. F.
figs. 12-13.
Assuming a specimen in the British Museum to be, as I fear it
is not, correctly named, the appendages, like the fly itself, are very
like dilecta. The clasps are longer, 1:13 mm. against 0:93 mm.,
the neck-like portion longer and straighter, 7. ¢., its sides are more
parallel and the spinous process is appreciably larger, bolder, and
less bent down.
I believe I have not seen a specimen of true coalita de Nicév.
These appendages are even more like those of rona (biagi) than
those of dilecta. If it be not coalita, it may be a form of rona,
or else a new species.
Text-fig. 87.
Named coalita in B.M. Coll. (probably erroneously). X 38.
22. cardia Felder, Sitz. Akad. Wiss. Wien, Math.-nat. Cl. xl.
p. 459 (1860).
Ihave felt some difficulty in discriminating this from dilecta
Moore, both in the superficial characters and in the appendages,
and am still doubtful whether they do not approach each other
within the limits of geographical races of one species—dilecta
being the continental, cardia the insular form; though such
a distinction does not quite agree with the localities of specimens
examined, a specimen from North Borneo being indistinguishable
452 DR. T. A. CHAPMAN ON THE SPECIES [Apr. 27,
from dilecta, and an almost typical dilecta was received from North
Luzon (Philippines).
Text-fig. 88.
cardia. X 43.
Text-fig. 89.
cardia. X 42.
1909.] OF THE LEPIDOPTEROUS GENUS LYCNOPSIS. 453
Text-fig. 90.
cardia (New Guinea). X 42.
23. dilecta Moore, P. Z. 8. 1879, p. 139.
This is a well marked species as occurring in continental India,
but I find in the Tring Museum unquestionable dilecta from
Borneo, Luzon, and Java.
The group to which it belongs is very puzzling, chiefly perhaps
because I am loth to recognise here unnamed species of which I
know little more than the actual specimen, and the appendages
of which differ by comparatively sight but apparently constant
differences.
There are some differences in Indian dilecta, due possibly to
slightly varied softening and pressing in mounting the specimens.
23 a, from Borneo, is exactly the same as some Indian specimens,
others from India are like 23.
236. A specimen from Perak has appendages closer to cosswa
than to anything else, the under side agrees closely with dilecta,
but the upper side has a darker blue and a dark border along the
hind margin.
23. A specimen labelled cossa (cossea 2) in the British Museum
belongs to this group, but the appendages differ from any named
species I have.
My material is too meagre to justify me in naming these two
forms.
454 DR. T. A. CHAPMAN ON THE SPECIES [Apr. 27,
Text-fig. 91.
dilecta. X43.
Text-fie, 92.
dilecta (Indian). X 41.
1909. ] OF THE LEPIDOPTEROUS GENUS LYCZNOPSIS. 455
Text-fig. 93.
dilecta (Upper Aroa River, Tring Coll.). x 41.
Labelled cossa, B.M. X 41.
456 DR. T. A. CHAPMAN ON THE SPECIES [Apr. 27,
24. cossea de Nicév. J. Bomb. N. H.S. ix. p. 271, pl. O. figs. 14, 15.
The dorsal process has a rougher (softer) inner and a smoother
(harder) outer portion. The clasps are large, very like those of
cardia or camene, but with the striking difference that the teeth
of the spinous process are directed distad instead of inwards.
Text-fig. 95.
cossea (B.M.). X 28.
25. lanka Moore, Ann. Mag. Nat. Hist. ser. 4, vol. xx. 1877, p. 342.
The spinous process of the clasp lies flat across the end, only the
point being free. Along the distal margin the secondary spines
are rather sharp and slender. No other species is quite near
to it.
26. rona Grose-Smith, Nov. Zool. i. p. 572, 1894 (no figure).
26a. biagi Bethune-Baker, is identical with this.
I have compared the type specimens and they agree perfectly,
and the coloration and marking of the under side is so different
from any other species that there can be no room for mistake.
The under side differs from other Lyczenopses, much as L. parr-
hasius does from H. argiades; 7.e. the markings, instead of being
dark on a pale ground, often black on white, are very pale
ochreous on a ground-colour of a not much paler tint.
My photograph of the appendages is from those of the type
specimen of biagi B.-B.
1909. ] OF THE LEPIDOPTEROUS GENUS LYCHNOPSIS. 457
Text-fig. 96.
lanka. X 42.
Text-fig. 97.
rona (biagi, type). X 4s.
Proc. Zoou. Soc.—1909, No. XXX. 30
458 DR. T, A. CHAPMAN ON THE SPECIES (Apr. 27
26 a. biagi Beth.-Baker, P. Z. S. 1908, p. 117, pl. vill. fig. 11.
There is a specimen in the British Museum labelled coalita
with appendages hardly distinguishable from those of biagi.
a
27. drucei Beth.-Baker, Ann. Mag. N. H. ser. 7, vol. xvii. p. 102;
P. Z. S. 1908, p. 117, pl. vin. fig. 16.
This well-marked species has appendages belonging to the
dilecta-group. The dorsal processes are larger than in dilecta and
the clasps broader, the spinous process much more developed,
and with teeth more obviously on its surfaces as well as its
margins.
28. akasa Horsf. Desc. Cat. Lep. E. I. Co. p. 67, pl. i. figs. 1, La
(1828).
This species is not likely on its general characters to be con-
founded with any other. The appendages show it to belong to a
rather large group of species in which the terminal spinous process
ig much reduced in size, but is still fairly developed, turned
inwards, but separated by a gap from the inner margin of the
end of the clasp.
29. camene de Nicév. J. Bomb. N. H.S. ix. p. 278, pl. O. fig. 22,
1895 (June ?1895).
29 a. selma H. H. Druce, P. Z. S. 1895, p. 573, pl. xxxii. fig. 10
(Oct. 1895).
Lam unable to see any difference in the appendages of these
two forms or indeed in the flies themselves.
There is a specimen in the Tring Collection from Mt. Dulangan,
Mindoro, that must be referred to this species. The clasps are
rather broader basally than in the typical specimens and the
lateral proximal process of the ring 1s smaller, shorter, and
sharper. These differences are, however, quite within the limits
of what may be ascribed to geographical variation.
A specimen labelled “‘ Perak—Curtis” differs rather more, and
Tam doubtful whether it may not be entitled to specific rank;
even if so, it is exceedingly close to camene. The clasps are
larger and bolder, and what may be called the neck occurs some-
what higher up. The fly itself is very dark on the upper side,
differing from camene much as lugra does from musina. The
under side is not definitely distinguishable, and this chiefly makes
me regard it as a geographical form. As camene dates from
Perak, it may really be that this is actually the typical form of
cumene.
A specimen from Sarawak in the Tring Museum is very small
and delicate (25 mm. expanse), with upper side very like dilecta,
except a broader border to fore wing, expanding towards costa.
1909. | OF THE LEPIDOPTEROUS GENUS LYCHNOPSIS. 459
Text-fig. 98.
druceit. X 42.
Text-fig. 99.
drucet. X 42.
460 DR. T, A, CHAPMAN ON THE SPECIES [ Apr. 27,
Text-fig. 100.
akasa. X 42.
Text-fig. 101.
camene. X 42.
1909.] OF THE LEPIDOPTEROUS GENUS LYCHNOPSIS. 461
Text-fig. 102.
(“ Perak—Curtis,” Tring Coll.) 48.
Text-fig. 103.
camene (selma, Mindoro, Tring Coll.). XX 43.
462 DR. T. A. CHAPMAN ON THE SPECIES [ Apr. 27,
Text-fig. 104.
camene (selma). X 43.
30. dilectissima H. H. Druce, P. Z. 8. 1895, p. 571, pl. xxxil.
figs. 2, 3.
The appendages are of the dilecta (cardia) type, smaller and
more delicate; length of clasp 0°82 (dilecta 0°93). The spinous
process more slender and sharper.
The clasp is also of a different form, broadest at the base,
whereas in dilecta it is broadest 2 of the way from the base and is
rounded in an elliptic line to an almost pointed base.
31. plata TW Eo WDruce EP. ZaSs 1895; pi at4, spl. scxxai-
figs. 8, 9.
The clasps have broad transverse ends without any teeth. It
is very similar to transpecta, with the considerable difference that
there are no large teeth on the inner margin as in that species.
There is a definite though small soft lower element to the dorsal
process.
Possibly I have not had true sonchus, the specimens examined
differing from plauta only in the deep yellowish tint of the under-
side; the appendages were identical with those ot plawta.
T have a suspicion (from figures and descriptions) that sonchus
and plauta are forms of one species.
1909. j OF THE LEPIDOPTEROUS GENUS LYCHNOPSIS. 463
Text-fig. 105,
dilectissima. X 38.
Text-fig. 106.
plauta. X 38.
464 DR. T. A. CHAPMAN ON THE SPECIES [Apr. 27,
32. sonchus H. H. Druce, P. Z. 8. 1896, p. 655, pl. xxix. fig. 4.
See plauta.
Text-fig. 107.
sonchus (or plauta). X 42.
33. melena Doherty, J. A. 8. B. lviil. p. 434, pl. xxiii. fig. 13.
This species belongs to the group in which the spicular process
of the clasp is of a length only half that of the extremity of the
clasp and lies down so close to the margin that it might conceiv-
ably be overlooked.
It is certainly very close to tenella. The clasps are, however,
decidedly longer and a good deal narrower.
34. tenella Miskin, Syn. Cat. Rhop. Austr. p. 63 (1891); Pr.
ihimns Soee Ni SaWe TOUS pe 144, 904 ios ial (Graeae
Waterhouse).
I have examined a specimen taken by Dr. G. G. C. Hodgson,
near Kuranda, between Naruba and Kuranda, at 1200 ft. elevation,
April 1899, and in his collection. As Cairns (at sea-level) is
within a score of miles, the locality is no doubt near if not the
same as that referred to by Miskin and Waterhouse as “ Cairns
District.” I assume my identification to be correct, as the under-
side markings agree exactly with Waterhouse’s somewhat dia-
grammatic figure. The blue has a somewhat greenish tone, not
alluded to in Waterhouse’s description.
The appendages are slightly smaller but otherwise quite
1909. | OF THE LEPIDOPTEROUS GENUS LYCNOPSIS. 465
Text-fig. 108.
melena, X 38.
Text-fig. 109.
tenella (placidula, Malay Peninsula, Tring Coll.). 44.
466 DR. 1. A. CHAPMAN ON THE SPECIES [ Apr. 27,
indistinguishable from those of L. owgarra (placidula) from New
Guinea.
1 entertain no doubt that they are races of one species. The
green tone of tlhe upper surface is a confirmation so far as it goes.
The differences are the large amount of white on the upper
surface of tenella, of which owgarra shows no indication in any of
the specimens I have seen but in one specimen of placidula there
are distinct white areas; and that on the under side the mark-
ings are much blacker in éenella, but the marginal markings have
only the marginal spots, at least the arched line that surrounds
them in placidula (and a majority of Lycenopsids) is only faintly
indicated.
Text-fig. 110.
tenella (placidula, Aroa River, Tring Coll.). X 45.
34a. Var. placidula H. H. Druce, P. Z. 8. 1895, p. 572, pl. xxxui
mes, Oy Me
34b. Var. owgarra Beth.-Baker, P. Z.S. 1908, p. 116, pl. viii.
mee, IT .
The clasps are very short and broad, and very broad at the
extremity, the spinous process is minute and fused to the margin
of the clasp along the dorsal third of the extremity.
It is very similar to melena and to singalensis. The clasp is
much broader than in melena, and singalensis is larger and has
the terminal half of the clasp distinctly narrowed.
1909.) OF THE LEPIDOPTEROUS GENUS LYCZNOPSIS. 467
Text-fig. 111.
tenella (owgarra, type). 42.
Text-fig. 112.
tenella (Australia). 45.
468 DR. T. A. CHAPMAN ON THE SPECIES | Apr. 27
It is no wise related to placida. placidula has a coloration
similar to that of placida; owgarra has a very green tint, looking
at first sight quite a different insect. In the Tring Collection is a
specimen, rather deep black-blue, but fairly typical placidula, from
Gunong Tjan, Malay Pens.; a very similar specimen, except that it
possesses white patches (like puspa) on all the wings, from Upper
Aroa Riv er, N.G.; a typical owgarra from the same locality, and
another owgarra from Obi Major.
This shows a very wide range for the species—Malay Peninsula,
Moluceas, Borneo, and Australia. The identity of the appendages
in specimens from these widely separated districts appears to
remove any suspicions one might entertain that sinrgalensis and
melena were varietally related to placidula; singalensis having
the clasps, &e., much larger, and melena@ much narrower,
wa
35. singalensis Felder, Verh. zool.-bot. Gesellsch. Wien, xviii.
p: 282.
The appendages are very like those of ¢enella ( placidula), the
spinous process being small and almost lost by being pressed down
on the upper angle of the extremity of the clasp, which ends in a
nearly transverse line. It differs by its larger size, 0°88 mm.
long, against 0°62, and by the ter ‘minal half being g very definitely
narrowed into a long neck-like portion. The whole clasp, and
especially the base, is ‘much broader than in melena.
De Nicéville seemed in much doubt as to this being a “ good ”
species (Butt. Ind. vol. 11. e 108), and Bingham sinks it as a var.
of huegelii. singalensis is, however, a very distinct species,
whilst hwegelit is a form oa or grolus.
In the Druce Collection there is a series of this species from two
localities in Sumatra. They are of a much darker, deeper blue
than those from Ceylon, with a strong tendency to a dark border
and to spots round the margin of the hind w ing; they rather
suggest very large, borderless puspa, such for example as one in
the Tri ing Coll. from Palawan.
In the Tring Collection is a very pale specimen from Java,
looking very much indeed like a large pale huegelit, with very
white fr inges on the hind wing and round the anal angle of fore
wing. One would hardly recognise this as the same species as
the Sumatran form without the indication afforded by the appen-
dages; the fore wing is even of a different form, being more
pointed. The under-side markings differ also, much as those of
strophis (or dilectissima) do from limbata.
De Nicéville refers to specimens from the Nilgiris ; an Indian
specimen in the British Museum labelled singalensis is oreas,
previously not known as Indian. singalensis is probably not a
continental form.
De Nicéville & Elwes refer specimens from Lombok to huegelii
(J. A. S. B. 1897, p. 695). As huegeliz almost certainly does not
extend to Lombok, these specimens in all probability are singa-
lensis. I have not, however, seen them.
1909.] OF THE LEPIDOPTEROUS GENUS LYCANOPSIS. 469
Text-fig. 113.
singalensis (Ceylon). % 33.
Text-fig. 114. _
singalensis (Sumatra, Druce Coll.). 42.
470 DR. T. A. CHAPMAN ON THE SPECIES [Apr. 27,
The figures of the appendages to same enlargement show the
great difference in size between those of singalensis and tenella ;
but the one of the former at a less amplification is hardly distin-
euishable from the latter. The species are probably closely related,
but are now quite distinct.
36. ceyx de Nicév. J. B. N. H.S. vii. p. 328, pl. H. tigs. 6, 7.
The appendages perhaps most resemble those of plauta; like it,
the clasps have gone beyond dilectissima and placidula in the
gradual dwindling of the spinous process, so that it is quite lost,
except a trifling greater chitinisation of the angle whence it has
disappeared. The clasps are very broad basally and joined
together for a rather longer distance than usual. The dorsal
processes have a well-marked, rather large, rounded elevation as
ventral accessory.
Text-fig. 115.
ceyx. X 40.
37. carne de Nicév. J. B. N. H. 8. ix. p. 274, pl. O. fig. 18.
T have not been able to examine this species, described from a.
unique specimen ; the figure suggests to me that it is a Sumatran
race of L. nedda.
38. albecerulea Moore, P. Z. 8. 1879, p. 139.
One of the most specialised and remarkable forms. The dorsal
processes have developed a terminal (not ventral as in dimbata, &c.)
1909. ] OF THE LEPIDOPTEROUS GENUS LYCHNOPSIS. 471
sharp point, and the clasps are very broad basally and narrow
rapidly to a sharp point, in form something like an inverted
pegtop. No other species has anything like either of these
peculiarities.
Text-fig. 116.
alhocerulea. X Al.
39: catreus de Nicév. J. B. N. H. 8. 1% p. 2/6, pl. O.
figs. 20, 21.
Butler questions this being a Cyaniris because of its neuration,
but his objection is, that its neuration is that which is typical of
Lycenopsis. I question it also because of the somewhat bizarre
form of the clasp, which is unlike anything else in the genus, and
because of the great development of the accessory ventral process
of the dorsal processes, and those dorsal processes themselves
possessing several terminal spines; these are small, but are
reminiscent of albocerulea, to which and to albidisca it seems
most nearly allied. The ventral accessories of the dorsal pro-
cesses have here nothing to do with the missing hooks, but
are rather the ventral portion of abdominal segment 10, and
are probably united by softer tissues across the middle line.
40. hersilia Leech, Butt. China, p. 319, pl. xxxi. fig. 16.
T have not heard of a ¢ of this species.
472 DR. T. A. CHAPMAN ON THE SPECIES [Apr. 27,
Text-fig. 117.
catreus. X 38.
Text-fig. 118.
Bothrinia nebulosa. X 41.
1909. ] OF THE LEPIDOPTEROUS GENUS LYCENOPSIS. 473
41. deliciosa Pagenstecher, Ent. Nach. xxii. p. 50; figured Abh.
Senck. Ges. xxiii. pl. xx. fig. 8.
I have not seen this insect and can say nothing beyond what
Dr. Butler remarks as to its doubtful claim to be a Lycenopsis.
BovruRrini, n. nom.
Lothria, amended from SLotria (Rondani), is in use for a
dipterous genus, therefore it is necessary to rename Bothria
(mihi, P. Z. 8. 1908, p. 677). I suggest Bothrinia.
42. nebulosa Leech, Ent. xxii. p. 43.
This species belongs to the Everids and to the same genus as
chennellit, viz. Bothrinia, and is not a Lycenopsid at all.
The appendages have the characteristic dorsal armature, the
two (usually) side pieces fused into one in the middle line (as in
Everes, Cupido, &e.), and the clasps with two branches of equal
and considerable length. In this case the outer or dorsal branch
has a flowing curve not dissimilar to that of the same branch in
Everes, but curiously both branches are much alike, each ending
in three obtuse but sharp spines.
The neuration (anastomosis of veins 11 and 12 of fore wing) is
also that of Everes, not of Lycenopsis.
The row of spots under the fore wing, like those of chennellii,
are in line; in Lycenopsis those above vein 6 are usually more or
less moved inwards. This is not a constant character as between
Everids and Lyczenopsids, but there is so much difference in this
respect that it may almost be so regarded.
43. chennelli is also an Everid (Lothrinia), not only not a
Lycenopsis, but not in the same group of Lycenids
(see Proc. Zool. Soc. 1908, p. 676).
ARTOPOETES, n. g.*
44, pryeri.
Whether on his own or what other authority Dr. Butler
(Ann. & Mag. Nat. Hist. vol. v. 1900, p. 442) places this species
in Lycenopsis, | do not know. I may say with Mr. Murray, who
first described it, “It is not closely allied to any species with
which I am acquainted.” The clasps are so divided that it has
the appearance of having two pairs of clasps, a condition I have
not noticed in any other (very few, no doubt) Lycenid I have
examined, though there is some resemblance to it in Tarucus nara.
* aptomovs—baker. (There is already a genus Artopoia.)
Proc. Zoou, Soc.—1909, No. XXXI. 3]
474 DR. T. A. CHAPMAN ON THE SPECIES [ Apr. 27,
Text-fig. 119.
Artopoetes pryeri. X15.
Text-fig. 120.
Megisba malaya. X 45.
1909. ] OF THE LEPIDOPTEROUS GENUS LYCANOPSIS. AT5
Text-fig. 121.
Neopithecops zalmora. X 45.
Text-fig. 122.
Castalius ethion. 45.
A76 SPECIES OF THE LEPIDOPTEROUS GENUS LyC&HNopsis. [Apr. 27,
It certainly does not belong to any genus the male appendages of
which I happen to have observed.
The width of the central vacancy in the dorsal armature seems
to ally it with the Lycenopsids, though the accessory hooks are
well developed; and I would propose for it a new genus,
Artopéetes.
45. Megisha malaya, might be a Lycenopsis.
46. Neopithecops zalmora.
AT. Castalius ethion.
Both these remind one somewhat of C. catreus.
Index of Species.
ACESIN Ale eeeR eee ena eer, Tarmac g erasure ene eam Sy
UIE HM nanntassapeeuetesanes ucts} | levettith 2. ten aene eee al\G
SMOWGHISED, — anaconacooascse LA INVTRICENAY, coe yaovooccovcs ccc 19
alboceruleoides ... .. 16 athe Satanbaccdekaunadatese MOG
albocerulea ............ 38 MySay (Wess cosee ta ae Ola,
APEACMMS ccoscaoooonacasces LG | IND Yas AY oc Sesame Mame aes 3a
Iberetavaree eee eel ove | TITEL s cco aondrdecaetdecs 45
PIAS Rec aaat cence: MOS | MaLcmmatale so. eeemeels
[OWA ocoooconan nae 2 | TNCIETTA conconsonse cave OS
bothrinoides ............ 16 | THMMGWIND eocnceacnconsscece 3
Cajal ae eee UYSISUNOSD, cco ssdgan conan 42
CREDITED. 550 cosc00c0000500¢ 29 | TOECIGEN sane cegnuemeeradecs © mnt)
CATAM Ai Etec eres | ONCAS Serer eee yr tae SLD,
Corio Masoeadhaontcancaenad 22) | OW GETEED, oso nspcnscossonoce 345
CALI Ae ee OT AVVO, ses -oeeccoee, UL
GHITWWS scossoscassdedoosa OLD | DOWISI® cacateccanasece Oy 7
COVKE Neiecomeneeaee Eee MOO ICO cocsoscvacshnoscke LOG
chennelliiee: eee PLACA eee cc
GHINGEENE) Gopoconasodtonoses G2 lala core 31
COME, ceonccecsoononocce VAL PLY CUI se aecn ees ed
GOMOSGVTE, sccsoncccaccean 1K pseudargiolus ......... 16
GOATS scoconsoossvece 4, IOWISIIE) oguormeobecomacucennn ls
(COPED, occccoctsoncevcccs Mtl puspargiolus............ 16
cyanescens\......-..------ 15 JDUSOMAWS coocagsnoeeoonn 1S
GUERTTITED nao noaconnanae 15 TMP LO mn ocaseceasccuccsees pala
GEING OS) con coo 500005 500000 Al OWEN. hogeeenemursreeosssan: 240
GME, accosooostcconccen BD} SDT cosbiacocsodenseacoces. LOG
dilectissima ............ 30 Sheliond ae eeeeeee eae 5
GbADL Sails Men neceanneememace. | Stim ae occ eG
UMN | gaasccdoacasdeacans: — CLz/ | Sing alensisuaeeeaeee 35
IDEVEANGHEIS soccenooncccona 183 SOnChusi yee
TACESMD, cocoscsocnocooeces 410) splendens ............... 15
IEE soconscconcaccooes IG | SHPO DANG ccccopoccorsece esc 20
THATOEANIEDE cooconcopongcce Alb | Gemelli ae eee eee 34
JP AMFSVTE; Loo sonnodcaccas LOD [DTG coceoceseacdeee US
dean eee 15 (HAITIAN cooccrscccenece 140)
ladloneye eres 16 | TWERGUIATE, sce ectocodneooe i
lad onid csi eee eee WAIQUOINED) Loo saecnacanann ace 16
lamblia demseeee reel TANLOROTR, socnasnecocosoncas ZAB
1909.] THE STRUCTURE OF THE CARNIVORE GALIDIA ELEGANS. 477
2. On some Points in the Structure of G'alidia elegans, and
on the Postcaval Vein in Carnivores. By Frank KE.
BepparD, M.A., F.R.S., F.Z.S., Prosector to the Society.
[Received March 18, 1909. |
(Text-figures 123-131.)
The death of a specimen of this Madagascar Viverrid on
February 21 last has enabled me to supplement my account of
Galidictis striata* with some notes upon its ally Galidia elegans,
of the anatomy of which there is at present but little knowledge.
The two genera are undoubtedly very closely allied, but never-
theless there are some differences which fully justify the generic
separation of the two, and which include facts of some little
interest as a contribution to the anatomy of the Viverride. I
have endeavoured to examine into all the matters which are known
to be of importance from a systematic point of view, and the
excellent condition of the specimen fortunately permitted a careful
study of some of the veins.
$ Katernal Characters.
The external characters of Galidia are for the most part known
and are referred to by Mivart in his account of the genera of
Ailuroidea 7. He observes that “ the anus does not seem to open
into any cutaneous depression.” I find that this statement, made
perhaps somewhat tentatively £, is perfectly correct. I also agree
with Mivart in finding no prescrotal glands.
Carpal vibrissz are present, as In so many other Carnivora $.
It will be noted that the absence of prescrotal glands dif-
ferentiates the genus Galidia from Galidictis, in which they are
present, but that both agree in the absence of a cutaneous pouch
into which the anus opens. The anus is large in Galidia, as is
also the case with Galidictis.
§$ Alimentary Canal.
y
In my paper upon Galidictis I have recorded the fact that in a
skull of Galidia elegans examined by me the first premolar was
present on both sides, there being thus—so far as this specimen
was concerned—no difference from the allied Hemigalidia. I
find, however, from an examination of the example of Galidia
elegans which forms the subject of the present communication to
the Society, that the tooth in question is absent, and from both
* “On sonie Points in the Structure of Galidictis striata,” P.Z.S. 1907, p. 803.
+ “On the Classification and Distribution of the Aluroidea,”’ P. Z. S. 1882,
p. 135.
{ It is queried in the definition of the subfamily Galidictine, loc. cit. p. 189.
§ Beddard, “ Observations upon the Carpal Vibrisse in Mammals,” P. Z.S. 1902,
vol. i. p. 127.
478 MR. F. E, BEDDARD ON THE STRUCTURE [Apr. 27,
sides of the jaw. The species therefore obviously varies in this
peculiarity. There is, moreover, a considerable diastema between
the canine and pm. 2, so that asmall pm. 1 which has not emerged
may be present in the jaw.
The ridges of the palate in Galidia elegans are reproduced in
the accompanying figure (text-fig. 123). There are six complete
semicircular ridges perfectly symmetrical, of which the first is not
so extensive laterally as the rest. This runs across between the
Text-fig. 123,
Palate of Galidia elegans.
two canines. The next ridge corresponds to the diastema, and
the remaining four to the three premolars. Thereafter follow
four half-ridges on each side, but the details will be best under-
stood by a reference to the figure referred to.
The tongue has been already partly described by Mivart *, who
has recorded a well-defined patch of spiny papillee upon the middle
of the anterior half of that organ. This, as I have already pointed
out, is exactly what is to be found in Galidictis. There is, how-
ever, no further description of that organ, which I show in the
accompanying drawing (text-fig. 124).
* P.Z.S. 1882, p. 498.
1909.] OF THE CARNIVORE GALIDIA ELEGANS. 479
The papille circumvallate, as shown in that drawing, are only
two, right and left, as in Vandinia, the Genet, Hyena striata,
Proteles*, three or even more (Arctictis) beg met with in other
Viverride. There is nothing remarkable in the size or shape of
these papille.
Text-fig. 124.
Tongue of Galidia elegans, dorsal view.
On the right are eight of the strong anterior papille more highly magnified.
The fungiform papille are scattered over the dorsum of the
tongue between the papille circumvallate and the anterior patch
of strong papille; they are chiefly massed in the actual middle
line of the tongue, and there is no narrow middle line free from
them as in the Civet. The arrangement of these papille seems,
* Loe. cit. p. 500. J can confirm Dr. Mivart in the case of Hyena striata, where
there are distinctly only two of these papille.
480 MR, F. E, BEDDARD ON THE STRUCTURE fApr. 27,
in fact, to be very like that of Vandinia. The anterior patch of
conical papille is like that of Galidictis, which has been figured
by myself *, and of about the same extent. A few of the papillee
are shown more highly magnified at the side. Their tips show a
tendency to become bifid and to be separated from the rest of the
papille. In these characters they are not unlike the conical
papille of Galidictis.
The frenum of the tongue has a free fold of mucous membrane
near its attachment to the floor of the mouth; but this fold does
not form a continuous fold anteriorly, but is prolonged into two
elongated processes on either side of the frenum, apparently dif-
fering from the Genet.
The stomach seems to me to be as described by Dr. Mivart t,
and I have nothing to add to his remarks.
The intestinal tract is quite typically Carnivorous, but there are
various details of structure to which it is necessary to refer in
comparing Galidia elegans with its allies, and especially with
Galidictis. The duodenum forms the usual C-shaped loop enclosing
a part of the pancreas, from the end of which (where the pancreas
bends up towards the liver parallel with the portal vein) arises the
ligamentum recto-duodenale as in other Carnivora. This ligament
has a curious relation to the hepato-caval ligament, which, if it be
more general than I suppose, is at least not universal. The latter
ligament, arising from the inner edge of the caudate lobe of the
liver, is attached as usual to the postcaval vein, but a considerable
portion of it strays on to the mesocolon and becomes continuous
with the recto-duodenal ligament, the two forming together a
semicircular fold of watch-pocket form (see text-fig. 125). I found
the same arrangement to be much more marked and symmetrical
in Vandinia binotata. At the opposite extreme is Procyon lotor,
in which the short ligamentum hepato-cavale is attached only to
the postcaval vein.
As is the case with other terrestrial Carnivora, the colon and
rectum form a very short tube, which is perfectly straight in
Galidia. It is decidedly shorter than in Galidictis, where I have
described a rudimentary transverse colon, and where, if the colon
were straightened out, it would reach forward as far as the
diaphragm. In Galidia the colon lies perfectly straight, and only
reaches up to the anterior end of the left kidney, at which point
the cecum arises. The total length of the colo-rectum is 4? inches,
something short, therefore, of that of Galidictis. J have made
notes on the form of the colon in a number of Viverrids, which I
may here compare with that tube in Galidia and Galidictis. In
Crossarchus fasciatus there is as well a developed transverse colon
as in Galidictis. Suricata tetradactyla agrees with the two last-
mentioned Aluroids. If the colon were fully straightened out,
the ceecum would be seen to arise a long way in front of the left
* PZ. 8. 1907, p. 806, text-fig. 210.
+ Mivart, P. Z.S. 1882, p. 499, fig. 3 B. t P.Z.S. 1882, Joc. cit.
1909. | OF THE CARNIVORE GALIDIA ELEGANS, 481
kidney. The species Herpestes smithii shows the same bending of
the colon over to the right side of the body. On the other hand,
in Paradoxurus hermaphroditus the colon is short and quite
straight, and there is no trace of a transverse colon, such as exists
in the types already referred to. The cecum of this animal, in
fact, lies pretty well in the middle of the body. In the closely-
allied Paradoxurus niger the conditions are the same, and the
cecum originates only just in front of the left kidney. Viverra
civetta is also quite like Paradoxurus. Genetta rubiginosa has this
Text-fig. 125.
Caudate lobe of liver, duodenum, and adjacent structures in Galidia elegans.
e. Colon. d. Duodenum, from which arises recto-duodenal ligament partly in-
serted upon mesocolon and partly continuous with hepato-caval ligament as
explained in text. AK. Right kidney. JZ. Caudate lobe of liver, from the
lowermost process of which arises hepato-caval ligament. p.c. Postcaval
vein.
peculiarity exaggerated, for the cecum arises from the colon con-
siderably behind the left kidney. While, therefore, these facts
seem to justify the separation of Viverrines and Herpestines
among the Viverride, they leave the affinities of the alleged sub-
family G‘alidictine doubtful; and it is remarkable that while
Galidictis is Viverrine in its scent-glands, it is Herpestine in its
colon, while, on the other hand, Galidia is Hevpestine in the
absence of scent-glands and Viverrine in the condition of the
colon, which is perhaps what is to be expected in a subfamily
482 MR. F. E, BEDDARD ON THE STRUCTURE [Apr. 27,
believed to be an ancient Viverrid assemblage. The relationship
of the colon to the left kidney is shown in the accompanying
figure (text-fig. 126). That drawing also illustrates the cecum
shown from both sides. It is necessary to exhibit the cecum on
both sides because of the differences in the czco-colic ligaments on
the two sides of the cecum. The cecum itself, independently of
its mesenteries, has been already figured by Dr. Mivart*, and
quite correctly except that it is repr resented as stra ighter than it
is in nature. This is, however, to be accounted for by the cutting
of the various ligaments which attach the cecum to adjoming
regions of the gut. It is clear, however, from this figure that
Galidia agrees sh Galidictis + in possessing a long and thus
Herpestine cecum. In my figure of the cecum of Culidictis I
Text-fig.'126.
Cxcum and adjacent structures of Galidia elegans viewed from both sides.
The left-hand figure shows the position of the cecum with regard to the kidney,
which is partly covered by the colon.
a. Median frenum of cecum. 6. Lateral cxco-colic ligaments.
ce. Lymphatic gland.
have contrasted this Viverrid with Herpestes fulvescens, in which
latter the frenum of the cecum—the anangious ligament binding
it to the ileum—is continuous up to the very top of the cecum.
In Galidictis this is not the case, and the actual tip of the cecum
is not thus attached ; the frenum does not run so far. Galidia
agrees entirely in this particular with Galidictis. I may take this
opportunity of mentioning that Suricata tetradactyla does not
agree in this full extension of the ligament to the very tip of the
cecum with Herpesies. I also pointed out in the paper referred
to that in addition to the median anangious frenum the cecum
* P.Z.S. 1882, p. 508, fig. 8 C. + P.Z.S. 1907, p. 805, text-fig. 211.
1909.] OF THE CARNIVORE GALIDIA ELEGANS. 483
of these animals is fixed to the gut by two minute lateral ligaments
which bear blood-vessels. These were particularly obvious in the
example of Galidia elegans which I have dissected, since they were
loaded with fat. Jt is noteworthy that the median membrane
being anangious had not received a coating of fat. ‘These mem-
branes, one on the left and one on the right side, are almost
microscopic, as is shown in my drawing of Galidictis*. Galidia
elegans shows an interesting difference, for in that Carnivore the
membrane on the right side is very much larger than that on the
left, and joins the median anangious frenum. ‘The figure (text-
fig. 126) also shows a lymphatic g gland imbedded in the right- hand
lateral mesentery, the presence of which appears to be very typical
in these Carnivora. I have already referred to its existence in
Galidictis striata and Genetta vulgaris. I have also found it in
Suricata. The left-hand mesentery is quite small.
Text-fig. 127.
Liver of Galidia elegans, abdominal surface.
c.a. Caudate lobe. g.b. Gall-bladder. 7.c. Left central lobe. 7.7. Left lateral lobe.
p.c. Posteaval vein. 7.c. Right central lobe. 7.J. Right lateral lobe.
s.p. Spigelian lobe.
The pancreas has the usual form of the figure 6 so general in
Carnivora. The circular portion of the gland lies within the
duodenal loop. The liver is represented from the abdominal
surface in the accompanying text-figure (text-fig. 127). It does
not differ greatly from that of Galidictis 7, with which it may be
compared.
* P.Z.S. 1907, p. 808, text-fig. 911 B.
ape fs Al 1907 p- 809, text-fig. 212.
484 MR. F, E. BEDDARD ON THE STRUCTURE [Apr. 27,
§ Brain.
I figure the brain of Galidia elegans in two aspects (text-fig. 128),
from the dorsal side and a lateral view, which together expose all
the fissures at any rate of one side. There is, however, no great
asymmetry in the brain of this Viverrine such as demands the
additional representation in the figure of the left side. On the
whole, the brain agrees very closely with that of Galidictis *, which
IT have already figured and described at some length, comparing it
with the brain of allied forms. Viewed from above, the general
outline of the brain does not present any marked differences from
that of Galidictis, and it is of about the same size.
Text-fig. 128.
LEE
Brain of Galidia elegans, dorsal view and lateral view.
a. Crucial suleus. 6. Precrucial sulcus. c. Lateral sulcus. d. Suprasylvian.
e. ? Ketolateral sulcus. jf, g. Kctosylvian fissures (see text). S. Sylvian.
The crucial sulcus is long, as it is in Galidictis (where I have
contrasted its length with the shortness of the same fissure in
Viverra civetta), and is also slightly asymmetrical in relation to
its fellow. The right-hand suleus is a little in advance of the left.
It is curious that while the same furrows in Galidictis are also
* P.Z.S. 1907, p. 814, text-fis. 216.
1909. ] OF THE CARNIVORE GALIDIA ELEGANS. 485
slightly asymmetrical, the asymmetry is precisely the reverse. In
this region the brain of Galidia shows an interesting feature not
shown in the brain of its ally Galidictis. On the left side of the
brain, and on the left only, as is shown in the text-figure, there is
what certainly appears to be a precrucial sulcus, such as is figured
by Dr. Elliot Smith in Viverra*, and is well known to be very
characteristic of the Arctoid division of the Carnivora +.
The lateral sulcus seems to be in all probability a conjoined
lateral and coronary sulcus, as I have given reasons for believing
to be the case with the lateral sulcus of Galidictis. There is no
great difference between these fissures in the two Carnivores now
under consideration. Posteriorly in Galidia the sulcus bends
inwards and then outwards, ending therefore in a semicircular
outline. It may well be that this is an indication of a vestige of
an ento-lateral sulcus such as is figured by Elliot Smith in Her-
pestes pulverulentus %.
There is, moreover, just a faint indication of an anterior pro-
longation to the inside of the lateral sulcus from the angle which
the latter makes when it bends outwards. I could find no definite
ansate sulcus in the brain of Galidia.
The suprasylvian sulcus ends posteriorly at the level of the
inward bending of the lateral fissure. Anteriorly also it does not
extend so far as does the lateral fissure, in both these points dif-
fering from Galidictis and from Herpestes as figured by Elliot
Smith.
The sylvian complex of fissures differed slightly on the two sides
of the body. The right side is represented in the annexed figure
(text-fig. 128). The fissure which I have lettered “ /” seems to me
to correspond with the postsylvian fissure of Herpestes, in which
case perhaps “S’” is not really the sylvian fissure proper, but an
anterior ectosylvian, while “g” is a posterior ectosylvian. On
the other hand, on the left side of the brain, while S retains the
same position and relations, the two fissures “f” and “g” form
only one fissure, the lower part of ‘“f” being absent, and the
upper part joining, or very nearly joining, ‘‘g.” The arrangement
here is, in fact, very like that of the left side of the brain of
Galidictis figured by myself. There is in favour of the former
interpretation of the furrows in question the fact that the furrow
which I have lettered “ S'” does not reach the rhinal furrow at
quite the right point of a true sylvian fissure. This, I think, will
be apparent after an inspection of my drawing. It seems, on the
whole, to be the most reasonable course to regard the true sylvian
fissure as not present, but to consider the fissure which I have
lettered S to correspond to the similarly-lettered furrow in
Dr. Elliot Smith’s figure of the Hyzna’s brain §, which he refers
to in the text as “the so-called ‘Sylvian fissure,’” and which is
followed by two fissures in the temporal region, as in Galidia.
* Cat. Mus. Roy. Coll. Surgeons, vol. i. ed. 2, 1902, p. 249, fig. 122.
+ Mivart, Journ. Linn. Soc., Zool. vol. xix. 1886.
{ Loe. cit. p. 254, fig. 625. § Loe. cit. p. 257, fig. 130.
486 MR. F, E. BEDDARD ON THE STRUCTURE [ Apr. 27,
§ The Postcaval Vein and its Branches in Galidia
and in some other Carnivora.
These veins (text-fig. 129 A) in the specimen of Galidia which I
dissected were turgid with blood, and therefore in a very favour-
able condition for anatomical study. I take this opportunity of
comparing them with those of other Carnivora which I have
dissected lately. The large postcaval vein is quite normal in
position—that is, it lies to the right of the aortaas infthe majority
Text-fig. 129.
ae
A
Postcaval vein and branches in Galidia elegans (A) and Nandinia
binotata (B).
j. Intercostal veins. 7. Lumbar veins. K. Kidneys. 7. Renal veins. sp, Spermatic
or ovarian veins. sz. Suprarenal veins. 2. Splenic branch of ovarian.
x’. Spleen.
of mammals. The renal veins are asymmetrical, that of the right
kidney entering the postcaval above the entrance of the left renal.
This asymmetry is, it will be perceived, the exact reverse of that
of Galidictis*, where the left renal enters the postcaval higher
* P. Z.S. 1907, p. 811, text-fig. 213.
1909. ] OF THE CARNIVORE GALIDIA ELEGANS. 487
up than the right. Hach renal vein, when close to the kidney,
divides into two branches, one lying above the other, which supply
respectively the dorsal and ventral side of each kidney, over the
surface of which they ramify, as is so usually the case among the
Aluroidea. This difference of symmetry in the renal vein is cor-
related with the fact that in Galidia the left suprarenal flows into
the postcaval independently of the left renal, whereas they join
in Galidictis. Furthermore, there are no additional renal veins
such as are present in G‘alidictis. The spermatic veins arise as
in Galidictis (and indeed most mammals), the left from the left
renal, the right from the postcaval vein considerably below the
entrance into the latter of the renal veins. Two intercostal
veins flow into the dorsal surface of the postcaval between the
left renal and the right spermatic vein, each of them, however,
very nearly corresponding in the point of entrance to the point
of entrance of those two veins. The postcaval receives lower
down two lumbar veins on each side, as in Galidictis. The exact
disposition of these veins in Galidia elegans is as follows :—On
the left side there are two veins, each of considerable width, which
lie respectively above and below the corresponding artery, which
therefore emerges from the aorta between them ; before passing
respectively anteriorly and posteriorly the two veins are joined by
a bridge which passes over the artery just before it divides. On
the right side the two lumbar veins unite just before opening into
the postcaval, and they are not united by a bridge distally.
In an example (text-fig. 130 B, p. 488) of Suricata tetradactyla
I found a closer agreement with Galidictis than with Galidia.
There were two renal veins on each side, arising and ending
precisely as I have figured them in Galidictis. The right ovarian
vein, though pouring its contents into the postcaval itself as in
most other Carnivora, reached that vein rather nearer to the left
renal than in either Galidictis or Galidia.
In another example of Stwricata (text-fig. 130 A) the conditions
were In many points the same, but in others a little different.
On the right side the renal veins were two, though ultimately
reaching the postcaval by a common trunk. The renal was single
on the left side, and this vein was more symmetrical with its
fellows of the right side than in the first specimen. There was
in this specimen an unusuai origin of the left ovarian vein. The
right was as in the last specimen. The suprarenal of the left
side was, as often in Carnivores, an important parietal vein to
begin with, of which the actual suprarenal twig is only an
affluent. Into the conjoined parietal and suprarenal, before it
flows into the renal vein, open two parallel ovarian veins (as in
Marsupials), which thus cross over the left renal and do not open
directly into it. The two lumbar veins posteriorly are single and
symmetrical in their points of opening into the posteaval.
A dissection of two examples of Crossarchus fasciatus showed
that the ovarian veins were disposed precisely as in Galidia and
Galidictis, The renal veins were single.
488 MR, F. E. BEDDARD ON THE STRUCTURE [Apr. 27,
In Herpestes smithi I found differences of detail which are
worthy of record. The right and left spermatic veins had the
same relations to the postcaval and left renal respectively as
in the other luroid Carnivora already dealt with. The position
Text-fig. 130.
Postcaval vein and branches in Swricata tetradactyla (A, B) and Cercoleptes
caudivolvulus (C).
Lettering as in text-fig. 129.
of the right vein was more like that which I have described here
in Suricata. 'The‘renal veins were single, and the left supra-
renal entered the postcaval independently of the left renal.
1909. ] OF THE CARNIVORE GALIDIA ELEGANS. 489
The lumbar veins were in some respects like those of Galidiu,
and in other respects differed. They resemble those of Galidia
in the fact that there are two veins on each side, and that while
those of the left side open separately into the postcaval those
of the right side join before doing so. There is, however, this
important difference between the two animals in respect of these
veins. In Herpestes the postcaval has already divided into the
two iliacs, into each of which therefore the lumbars really open.
In Galidia the vein is a single vein at the points of entry of the
lumbars, and does not divide until later. It shouid also be
mentioned that the renals are asymmetrical with the asymmetry
of Galidia. With this is associated the separate entry into
the postcaval of the left suprarenal already mentioned. I say
“associated,” since in one specimen of Crossarchus where the
left renal and the left suprarenal entered the postcaval sepa-
rately, there was an asymmetry between the two renals; whereas
in a second example, where the left renal and suprarenal entered
the postcaval together, it was at a point exactly opposite to the
entrance of the right renal.
I have also collected some notes on Viverrine luroids, and
have already figured in Genetta vulgaris * the veins with which
IT am now dealing. In Paradoxurus hermaphroditus the two
renal veins were asymmetrical, the right entering the postcaval
above the left. The spermatic veins showed an exaggeration of
the condition described above in Suricata. The left spermatic
vein as usual enters the left renal; the right spermatic vein, as
is also quite usual with Carnivora, enters the postcaval directly
but very far forward—on a level, in fact, with the entrance
of the left renal. The two lumbar veins are, as in Genetta
vulgaris, single veins on each side. They enter the postcaval
asymmetrically, and are asymmetrical in two ways. In the
first place, the left vein enters rather more anteriorly than
the right. In the second place, the right vein enters the post-
caval after it has bifurcated to form the iliacs. The left vein
enters the postcaval in its undivided region. It is to be noted
that the corresponding arteries are also correspondingly asym-
metrical in the position of their exits from the aorta. I found
two examples of the African Civet Cat (Viverra civetta), both of
them males, to differ slightly im the branches of the post-
caval. In one the veins were exactly as I have described them
in Paradoxurus hermaphroditus, except that the lumbars were
quite symmetrical in position and that they both entered the
postcaval before its bifurcation. In the other the right spermatic
vein entered the postcaval some way behind the renals, and the
lumbars, though symmetrical, were two on each side.
Of Nandinia binotata (text-fig. 129 B, p. 486) I have examined
some of the ves with which I am concerned in the present
communication in two individuals, both males. In one of these the
* P.Z.S. 1907, p. 812, text-fig. 214,
Proc. Zoou. Soc.—1909, No. XX XII. 32
490 MR. F, E. BEDDARD ON THE STRUCTURE [Apr. 27,
two renals were pretty symmetrical as to their point of entry
into the postcaval. The left spermatic entered the left renal as
usual, and the right spermatic entered the postcaval as usual some
way further down. ‘The second example showed a very remark-
able variation—for the condition which I am about to describe
must be, I think, a variation, though I possess no positive notes
or sketches of anything similar in the first-described example *.
In this specimen the suprarenal, renal, and spermatic veins were
all separate from each other up to their entry into the postcaval
vein, which they entered in the order named. The spermatic
vein, however, was an unusually thick vein (for a spermatic vein),
and bending backwards ran for some little way parallel to the post-
caval, thus suggesting a persistent left cardinal vein. Presently
it divided into two, the smaller continuing in the same straight
line as a quite slender spermatic vein. The larger trunk bent
round to the left, and after giving off some branches to the
omentum ended in the spleen. There can be no possible doubt
about the course of this vein: it was full of blood, and could be
followed throughout with perfect ease. Furthermore, I do not
think that any affluent from the spleen reached the portal system.
All the blood thence derived flowed along the vein which I have
just described. This, again, perhaps favours the supposition of
the persistence of a left cardinal vein. On the other hand, this
view is not supported by the position of the orifices of the lumbar
veins. ‘These veins, in fact, of both sides open into the post-
caval. They are symmetrical, and there are two veins on each
side, as in many of the other Carnivora already described in the
present communication. Moreover, on the left side, and the left
side only, the two veins were joined by an anastomosis as in
Galidia.
I shall now give an account of these same veins in a number of
genera of Arctoid Carnivora.
In Procyon lotor the renals arose symmetrically. The left
infrarenal entered the postcaval independently of the left renal.
The right ovarian vein entered the postcaval some way below
the renals, into the left vein of which pair the left ovarian vein
poured its contents. There is thus no difference from the
arrangement met with in the majority of diluroid Carnivores,
except in the symmetry of the renals, which is not a very common
character of the Ailuroidea.
In Cercoleptes caudivolvulus (text-fig. 130 C, p. 488) there was
the same symmetry of the renals ; while the position of the ovarian
veins was also the same as in Procyon. The left suprarenal
joined the left renal just before the latter debouches into the
posteaval. The lumbar veins were single and slightly asymme-
trical, the left being in advance of the right. Hach lumbar gives
off a marked superficial branch running anteriorly dorsal to the
* In a specimen dissected subsequently (26. V. 9) there was no splenic yein
opening into the postcaval.
0) i OF THE CARNIVORE GALIDIA ELEGANS. 49]
kidney. Between the renals and the lumbars the postcaval
received two intercostal veins.
T have examined four individuals of /etonyx capensis—three of
them females and one male—which showed certain individual
differences. In one of them (text-fig. 131 B) the postcaval was
double postrenally, after the fashion of the Armadillos and
Anteaters *. The division was prolonged upwards to just ante-
riorly to the entrance of the left renal vein. The right rena
vein entered the postcaval higher up, where it was a single vein.
Text-fig. 131.
A
JS.
Ne
aay Ze
Posteaval vein and branches in Mellivora signata (A), Ictonyx capensis (B), and
Mephitis mephitica (C).
Lettering as in text-fig. 129.
The ovarian veins were symmetrical and arose each from its own
separate postcaval. This was also the case with the lumbar veins,
which were single veins. In the second female the conditions
were of the more “normal” Carnivorous type. The renals were
symmetrical, the postcaval was not divided, and the ovarian veins
had the very asymmetrical origin found in nearly all Carnivora.
* Hochstetter, Morph. Jahrb. xx. 1893.
32*
492 MR. F. E. BEDDARD ON THE STRUCTURE [Apr. 27,
In a third, also a female, the renal veins were asymmetrical,
but it was otherwise like the second specimen just described.
The fourth example, a male, was just like the last specimen in all
the veins the arrangement of which I studied.
I have preserved the skin of the “ abnormal” example and am
able to compare it with the skin of the second specimen which I
have described and with two others. There are, however, no
reasons for regarding the specimen as of a different species or
race. It is certainly a good deal smaller, which may of course
ultimately prove to be a racial or even specific character. But I
am unable to press this conclusion at present.
Three examples of J/eles meles differed among themselves. In
one there was what I term the typicalarrangement. In a second
the left ovarian vein, instead of opening on to the renal, had
moved down and debouched directly into the postcaval, but
above the entrance of the right ovarian vein. In this example,
moreover, the renals were symmetrical. In a third example the
renals were also symmetrical. But in this specimen the orifice
of the right ovarian vein had moved anteriorly and had come to
open into the postcaval in common with the left ovarian.
In a female example of Mephitis mephitica (text-fig. 131 C,
p. 491) I found a condition of the postcaval and its affluent veins
almost exactly like that of the first specimen of Jctonya of the
series just described in one important particular, but differing
in some other details. The point of resemblance between these
in other respects closely allied Mustelines was the division of the
posteaval vein postrenally. The division, however, in Mephitis 1s
more marked than even in Jetonyx. It extends, in fact, further
forward beyond the entrance of the left renal in Mephitis. Thus
the left renal pours its blood into the left half of the postcaval,
while the right renal communicates with the undivided anterior
region of the postcaval. There are, however, two renals on the
right side, though only one on the left. The ovarian veins arise
symmetrically in relation to their position in the body, but
asymmetrically in relation to the veins with which they are
connected. The left-hand vein is as usual a branch of the left
renal vein; the vein of the right opens as usual into the post-
caval, but at a point exactly opposite to the entrance of the left
renal and therefore from the right-hand subdivision of the post-
caval. Jt is noteworthy that here, as in some other cases, the
right ovarian vein is not a lateral affluent of the postcaval ; it flows
into that vein on its ventral surface.
In Bassaricyon alleni, the veins of which I have not referred to
in my account of the general anatomy of this animal *, the dissec-
tion of a second specimen enables me to describe the disposition
of the posteaval and its branches. The two renals are perfectly
symmetrical and the ovarian veins are disposed in the usual asym-
metrical fashion. The lumbar veins are double, but unite just
* P,Z.S. 1900, p. 661.
1909. | OF THE CARNIVORE GALIDIA ELEGANS. 493
before entering the single postcaval; neither pair is connected by
an anastomosis.
In three examples of Vasua rufa the branches of the postcaval
were practically identical. In all of them the asymmetry of the
renals was unusual, the left lying more anteriorly than the right.
From the left arises, as usual, the left ovarian (or spermatic), while:
the right-hand vein enters the postcaval below this point.
In the Otter (Lutra) the renals were quite symmetrical and the
ovarian veins quite as usual.
In Helictis personata the left ovarian vein was connected quite
as usual with the left renal vein. But before entering it the
ovarian vein received a small branch from the body- -wall in the
region of the kidney. This vein appears to me to be very possibly
the equivalent of the second lower and smaller renal veins which
I have figured in Galidictis. The right ovarian vein, as usual,
communicates directly with the postcaval.
In Ursus syriacus the ovarian veins presented an arrange-
ment which [ have not found in other Carnivora. Both were
directly connected with the corresponding renal vein. Nor were
the renal veins symmetrical, as perhaps might be inferred from
this fact. The right was in advance of the left, as is so commonly
the case not only in the Order of the Carnivora but in Mammals.
generally.
It is hardly possible to extract from the foregoing series of
facts any very plain cut-and-dried definitions of the several
groups of Carnivora derived from the characters of the postcaval
vein, such as is possible for example with the Marsupials. One
can only point out tendencies to the development of a particular
type in one group and of another type in another group of
Carnivores. Thus itis the Arctoid Carnivora, and possibly chiefly
the Musteline division of that group, in which the double post-
renal postcaval vein is to be met with*. The Arctoid genera, in
their wide distribution, nearly plantigrade feet, only at most
slightly specialised carnassial teeth, and with their non-retractile
claws, lie at a lower level than the A#luroidea. This result may
be perhaps taken into consideration along with the condition of
the postrenal section of the postcaval. In the same direction also
points the more usual symmetry in the position of the renal veins ;
these very frequently, more frequently than in the A¢]uroidea,
open into the postcaval opposite to each other. Again, itis more
common in this subdivision of the Carnivora for asymmetry to be
shown in the position of the spermatic or ovarian veins. No
* T am indebted to Mr. Burne for the information that in the Walrus the post-
caval vein is double from the level of the renal veins. This observation was made
upon the viscera of a young Walrus which died in the Society’s Gardens, and was
acquired by the Royal College of Surgeons. This condition in the Walrus is quite
possibly normal, for Mr. Burne has directed my attention to the fact that Dr. Murie
(Trans. Zool. Soc. vii. p. 431) found the vein double in the individual dissected by
himself. This doubling of the postcaval behind the kidneys is not, however,
distinctive of the Pinnipedia, for Murie figures (Trans. Zool. Soc. viii. p. 546,
woodcut fig. 4) a single postcava in the Patagonian Sea-Lion.
494 MR. F. E, BEDDARD ON THE STRUCTURE | Apr. 27,
absolute line of distinction can, it is true, be drawn between the
Arctoidea and the Ailuroidea in the arrangement of these veins.
But whereas among the Afluroidea only a few forms (e. g.,
Suricata, Herpestes smithi) can be mentioned in which the veins
are approximately symmetrical, there are more Arctoidea in which
this symmetry occurs, viz. in the species Mephitis mephitica, [ctonyx
capensis, Ursus syriacus. Moreover, in Ursus an absolutely
‘‘Marsupial ” disposition of the spermatic veins is to be met with,
where both are connected with the renal veins. Finally, the
lumbar veins are more usually restricted to a single vein upon
each side of the body in the Arctoidea; while they are prevalently
double in the Ailuroidea. That this feature is more archaic than
that which characterises the Alluroidea can hardly be said. I
merely dwell upon the fact that there is a tendency to the
establishing of this difference between the two groups. But in
dwelling upon these facts it must not be forgotten that some
Arctoidea are apparently exactly like some Ailuroids in the dis-
position of these veins.
Doubtless it cannot be said that the comparisons and contrasts
amount to a great deal. But they have at least an interest in
connection with the variations in the arrangement of the postcaval
and its branches in the Common Cat, “ Felis domestica.” The
veins of this animal have been lately studied in so large a number
of specimens as 605*. This large number probably gives some
real idea of the range of variations in these veins. In no less
than 23 examples the author found in the posterior region a
double postcaval, such as I have described in Jetonyx and
Mephitis. In 7 there was a left postcaval, instead of the normal
Kutherian right, In others, again, the spermatic veins, instead
of their normal asymmetry which is certainly the prevalent dis-
position in the Carnivora, as I have pointed out, both entered
the postcaval directly.
$ Other Blood-vessels.
The cerebral arteries of Galidia elegans are like those of other
Carnivora, differing only from those of certain forms in details.
There is at the posterior end of the medulla a strongly marked
rhomboid of arteries, as in many other Carnivora. This is asso-
ciated with a large anterior spinal artery and vertebral arteries.
The latter are single where they join the lateral angles of this
rhomboid, and not double as they are in Helictis personata *.
The middle cerebellar arteries are two on one side and one on the
other (left); but the latter divides into two very shortly after its
emergence from the basilar artery. It is to be noted that these
* W. Darrach, “ Variations in the Postcava and its Tributaries, as observed in
605 Examples of the Domestic Cat,’ Amer. Journ. Anat. vol. vi. 1907, Anat. Rec.
p. 30.
+ Beddard, “On the Anatomy of Helictis personata,” P. Z.S. 1905, p. 22,
text-fig. 8.
1909. ] OF THE CARNIVORE GALIDIA ELEGANS.
arteries arise behind the sixth nerves. The anterior cerebellar
arteries arise very shortly after the bifurcation of the basilar.
Excepting for the fact that the cerebellar arteries are strictly
paired, I found the arteries of the Common Otter precisely as has
been just described for Galidia.
Dr. Tandler* has figured the cerebral arterial system of the
Seal, and has represented the anterior spinal as of large size, as 1s
the case with many Carnivora including those to which I have
referred in the present communication. In a specimen of the
Common Seal which I dissected some time since this was not the
case. ‘The anterior spinal was very slender, and communicated
with the two very stout vertebrals by two very slender branches.
The middle cerebral arter My is single, and lies, as I have mentioned
in Galidia, behind the origin of the sixth nerve tT. The cirele of
Willis is joined anterior ly. by the anterior commennioatind artery.
I am unfortunately not quite certain about this part of the
cerebral arterial system in G'alidia; but I believe it to be like
that of Helictis figured by myself ¢.
The system of cerebral arteries in an example of Cryptoprocta
Jerox may be compared with that of the types already described
by others $ as well as by myself. The first remarkable fact is the
asymmetry of the arteries at the end of the medulla, which, how-
ever’, is very possibly an individual peculiarity, as the asymmetry
in the cerebral arteries of yraa descrihed by myself || proved to be
later §. Both the anterior spinal and the vertebral arteries are
stout. Although the main paired middle cerebellar artery lies
behind the sixth nerve, there is a slender branch on the right side
which lies in front of it. The anterior cerebellar artery hes just
behind the bifureation of the basilar. The arrangement of the
anterior communicating artery and of the anterior cerebrals
generally is very much as I have figured in Viverra civetta**.
I do not think that the cerebral arterial system of Zyana crocuta
has been deseribed. The specimen which I have had the oppor-
tunity of examining showed some differences from other Carnivora.
Beginning as before with the posterior end of this arterial system,
it is to be noted that the vertebral arteries are large—which is a
general character of the group. On the other hema, there is no
definite rhomboid-shaped arterial area at this point—z. ¢., at the
posterior end of the medulla—which is so characteristic of Carni-
vores. There are, in fact, two anterior spinal arteries, smaller than
the vertebral, which run back at least for some little way without
joining. I have no evidence, in fact, that they do ever join; for
* “ Zur vergl. Anatomie der Kopfarterien bei den Mammalia,” Denkschr. k. Akad.
Wiss. Wien, Bd. Ixvii. 1899.
+ I have already (P.Z.S. 1904, vol. i. p. 191) directed attention to the varying
position of these Bunnies with reference to the sixth nerve.
3 12 Z. S. 1905, p p.- 22, text- fig. 8
§ See also Bertha de Vriese, “Sur la Signification morphologique des Artéres
cérébrales,” Arch. d. Biol. xxi. 1904, p. 357.
|| P.Z.S. 1904, vol. i. p. 187, text-fig. 18. { P. Z.S. 1909, p. 166.
a* PZ... 1904, vol. 1. p. 194, text-fig. 21 B.
496 - MR. F. E. BEDDARD ON THE [Apr. 27,
the brain was severed from the spinal cord some little way behind
the medulla before I commenced its study. Into the circle of
Willis the carotids enter rather far back; and in front of them,
but still behind the sylvian artery, there was an ophthalmic
artery, as I presume it to be, on the right side only: I found no
trace of one on the left. A remarkable feature about the circle of
Willis of this Hyzena is the great length of the anterior commu-
nicating artery, which lay in a loose circle posteriorly. Generally
speaking, this artery is short and straight.
The iliac arteries in the specimen of Galidia were not sym-
metrical, as they are stated to be by Mivart * in other diluroidea.
The two larger external iliacs are produced by the terminal final
bifureation of the aorta. From the left of these arises a trunk
which continues for a short distance in the same straight lne as
the aorta and then divides into the two internal iliacs. It is
perhaps worth while mentioning that I found an asymmetry the
precise converse of this in a Squirrel (Sciwrus prevosti). In this
animal the aorta ended in the same way, by dividing to form the
two external iliacs. From the right of these arose a branch
which shortly divided to form the two internal iliacs.
3. On the Posteaval Vein and its Branches in certain
Mammals. By Frank KE. Bepparp, M.A., F.R.S.,
F.Z.8., Prosector to the Society.
[Received April 8, 1909. }
(Text-figures 132-139.)
In studying the venous system of a large number of Mammals
at the Prosectorium at the Zoological Society’s Gardens I have
relied entirely upon naturally injected veins. And indeed veins
turgid with their own blood are better for this study than arti-
ficial injections. For, in the first place, error is absolutely elimi-
nated, inasmuch as no injection-fluid can escape and give rise to
apparent branches which have no existence; and, secondly, it is
always possible to press upon the contained blood, and thus
ascertain the reality of anastomoses, which can be readily missed
in a defective artificial injection. Moreover, in animals which
have died with their veins full of blood, minute branches which
might be missed in other specimens, and which might not be
reached by an artificial injection, are plainly revealed. I have,
therefore, in the present communication only dealt with those
out of the many specimens which I have dissected during many
past years that were in a good condition for this particular
study, and have rejected the data derived from anzemic examples.
Thus I have every reason to believe that the facts which I now
bring before the Society are accurate. It is now well known
* P. Z.S. 1882, p. 515.
1909. | POSTCAVAL VEIN IN MAMMALS. AQT
that the venous system of Mammals, even as concerns the large
trunks, is subject to considerable variation. It is, therefore,
not without usefulness merely to record the facts alone as a con-
tribution to this department of anatomical study. I have, how-
ever, referred to previous work upon the subject, and have
endeavoured to summarise the present state of our knowledge
upon the posteaval veins of Mammals and their branches. The
important work of Hochstetter *, and more recently of McClure?
and others, upon the venous systems of Edentates and Marsupials
has attracted a great deal of attention to the venous system, and
has been productive of definite ideas as to the arrangement of
the several trunks in these animals. Jam able to confirm, and
in some respects to slightly extend, the work of these writers.
In making observations and in deducing conclusions from the
arrangement of the veins in Mammals it is requisite to bear in
mind the variability of these vessels, which appears to be greater
than that of the chief arterial trunks. Recently the variability
of the postcaval and its branches has been studied by McClure
in the Virginian Opossum (Didelphys marsupialis) £, by Wilhani
Darrach in the common domestic cat§, and by H. von W. Schulte
in various Marsupials||, while I myself have recorded4{[ some
variations among the Carnivora. J am able in the present com-
munication to note some variations as well as normal arrange-
ments. But inasmuch as variation occurs so frequently in so
fixed a type as the Carnivore Felis, a specialised race of a
specialised group, it is probable that variation occurs pretty well
everywhere. But in the centre of this variation lies a mean
which can be derived from the study of many examples.
§ The Postcaval Trunk and its Branches in various Orders
of Mammals.
The Marsupials show a very constant condition in that the
posteava lies medianly ventral to the aorta, so that on dissection
the aorta is not seen, being completely covered by the postcava.
The discovery of this very nearly universal characteristic of
Marsupials we owe, as Hochstetter has pointed out**, to Owen TT.
After examining Macropus giganteus, M. bennetti, Phascolomys
wombat, Phalangista vulpina, Didelphys lanigera, D. pusilla, Phas-
cologale penicillata, Belidews sciureus, Cuscus sp., Hypsiprymnus
sp., and two pouch-young of Petaurus taguanoides, Hochstetter
found that this generalisation held good for all those species with
* “Beitrage zur Entwickelungsgeschichte des Venensystems der Amnioten,’’
Morph. J.B. 1893; and “ Monotremen und Marsupialier,’ in Semon’s ‘ Forschungs-
reisen in Australien,’ Bd. ii. Lief. 3, 1896.
+ Amer. Journ. Anat. vol. ii. 1903, and vol. vy. 1906.
{ Loe. cit.
§ “Variations in the Postcava....in 605 Examples of the Domestic Cat,”
Amer. Journ. Anat. vol. vi. 1907, Anat. Rec. p. 30.
|| Ibid. p. 34.
{| “On the Anatomy of Galidia,” P. Z.S. 1909, p. 486.
** Toc, cit. p. 626. +t ‘The Anatomy of Vertebrates,’ vol. iil. p. 552.
498 MR. F, E, BEDDARD ON THE [ Apr. 27,
the exception of the last-named, Petawrus taguanoides. In that
species he found that the postcava lay to the right ‘‘as in the
Cat.” My own observations quite bear out those-of Hochstetter,
to whose list Iam able to add a few species dissected by myself
which that anatomist had not the opportunity of examining.
Tt have never met with the condition which characterises Petawrus
taguanoides in any Marsupial.
In addition to these types, McClure, as has already been
mentioned, has dissected many examples of Didelphys marsupialis,
as well as the Wombat and Petrogale sp.*, while Parsons has
reported upen Petrogale xanthopust, and Parsons and Windle
upon JJacropus rufust. The figure of Petrogale given by
McClare$ quite agrees with that of Macropus bennetti illustrating
Hochstetter’s remarks ||, In neither of these figures is the
anterior spermatic veins represented, though McClure states that
they are present, as also in the Wombat. They are furthermore
represented by the last-mentioned anatomist in Didelphys4, and
the anastomosis between the anterior and posterior spermatic
veins clearly shown. Inasmuch as McClure has remarked that
in the Wombat the spermatic veins also open in the neighbourhood
of the kidneys, as well as into the postcaval trunk, I do not quite
understand his saying, in a later paper, that “In a number of
adult Australian Marsupials, however, the spermatics do not open
into the postcava, as in Didelphys, but open into it at the base
of the renal veins, as in Phascolomys wombat, or into the renal
veins themselves, as in Votoryctes typhlops,” as observed by Miss
Sweet**. It would appear, however, that McClure does not allow
the value of a spermatic vein to a vessel often slender which does
pass between the gonads and the renal vein; for he writes
thus of Didelphys: ‘In none of the adults examined did the
spermatics open into the renals, although an anastomosis between
the latter and the spermatics was invariably TY present, on each
side, in the form of a small vein which followed the ureter.” As
a matter of fact, I have myself generally found two such small
veins which were frequently large veins fully as important as
what I term the posterior spermatic veins, which open behind
them into the postcaval itself. Nor, indeed, is any difference of
size shown by McClure in the plate{{ which illustrates the main
venous trunks (as well as the arterial) of Didelphys virginiana.
In a female Zrichosurus vulpecula 1 found that the postcaval
and its various branches were arranged as follows :—The asym-
metrical renal veins not only lacked symmetry in their plan of
opening into the postcaval trunk, but also in their number, for
* Amer. Journ. Anat. vol. ii. 1903, p. 388. + P. Z. S. 1896, p. 683.
~ J. Anat. Phys. vol. xxxii. 1898, p.119. This paper deals mostly with osteology,
muscles, and viscera. But the authors mention the ventral position of postcaval.
§ Loe. cit. p. 388, fig. v1. || Loe. cit. p. 626, fig. 13.
@ Amer. Journ. Anat. vol. v. 1906, p. 199.
** Proc. Roy. Soc. Victoria, vol. xvii. 1904, pt. 1.
++ Italics Dr. McClure’s.
te Amer. Journ. Anat. vol. v. 1906, pl. i.
1909. ] POSTCAVAL VEIN IN MAMMALS. 499
on the right there were two of these veins of equal and con-
siderable size. On the left only a single renal vein debouched
into the postcava. As is usual (? universal) among Marsupials
both renal veins give off an ovarian vein, in addition to that pair
which flow directly into the postcaval. On the right side a single
Text-fig. 132.
Posteaval vein and its branches in Tvrichosurus vulpecula (A) and
Trichosurus fuliginosus (B).
i. Intercostal veins. K. Kidney. 7. Lumbar veins. +. Renal veins.
sp. Spermatic or ovarian veins.
vein opened into the lower of the two renal veins; at a little
distance from its point of opening into the renal vein this ovarian
vein divided into two. Of these the outer branch opened into
the lumbar parietal some little distance away from the opening
500 MR. F, E. BEDDARD ON THE [ Apr. 27,
of the latter into the postcaval. The inner vessel, running of
course parallel with the outer branch, crossed over the lumbar
and reached the ovary. There was no junction between this mi
and the main ovarian trunk, which I shall describe directly a
opening straight into the posteaval, except perhaps quite distally,
where the smaller branches of both these veins may anastomose
on and about the ovary.
On the left side the veins in question were somewhat different.
There are, however, two of them, as on the right side. Anteriorly
these two veins cross and anastomose with each other more than
once, and one of them appears to be connected directly with the
kidney of the left side. Posteriorly the two vessels are again
united by a short transverse trunk on a level with the posterior
ovarian veins. Both these vessels appear to reach the ovary.
With the left lumbar parietal vein is connected a forwardly
running branch, as on the right side. Jt occupies an entirely
corresponding position, but does not form either of the ovarian
veins just described. It gradually dies away anteriorly. Between
the left renal vein and the lumbars already spoken of are four
intercostal veins. On a level with the last of these and rather
asymmetrically (the left being in front of the right) enter the
posterior ovarian veins.
Of a few other Marsupials I have less complete notes upon
these various veins. Ina specimen of Zvrichosurus fuliginosus*
the two renals were very asymmetrical, the left being much below
the right in its point of entry into the postcaval ; the two anterior
sper matic veins were present, and the left certainly, and the right
probably, flowed into each renal. Whether they segues
with the posterior spermatics I am not certain, but I think that
they did not. The latter veins were also asymmetrical, the right
opening into the postcaval below the left.
A second example of TZrichosurus fuliginosus showed some
additional features of which I had not made notes in the first
example. The asymmetry of the posterior ovarian veins seems
to me to be explained by this specimen. When the right vein is
earefully examined it is seen not to enter the postcaval, where it
appears to enter, i.e. just opposite to the entry of the left ovarian
vein. From this point it runs forwards, closely adhering to the
posteaval vein until it reaches the next intercostal vein, in common
with which it opens into the postcaval. Whether there i is or is
not a connection also with the intercostal lying next behind the
one just referred to—and I am inclined to think that there is
not—we have here, as I believe, a retention of a more anterior
section of the postcardinal than is retained on the left side. This
appears to be shown by the course of the vein strictly parallel to
and in close contact with the postcaval (i. e. the right subcardinal,
* In all the species which I describe here the postcaval lies below (7. e. accu-
rately above, en a dissection from the ventral surface) the aorta. This fact, clearly
unusual among Marsupials, was first discovered by Owen, as both Hochstetter and
McClure have pointed out in their memoirs.
1909. | POSTCAVAL VEIN IN MAMMALS. 501
or both cardinal collaterals*) and its connection with a vein of
the body-wall.
On the left side two vessels run, one on each side of the ureter
to the renal vein of that side into which they opened after fusing
into one vein. Lower down, these vessels were connected by
several cross-anastomoses. The outer of the two, which is the
real anterior ovarian vein, anastomoses with the posterior ovarian
vein only just in front of the ovary. The inner and more slender
of the two vessels could be distinctly seen to pass dorsally of the
posterior ovarian, and to follow the ureter to the bladder. On
the right side the outer (ovarian) vessel did not fuse with the
posterior ovarian, but passed dorsally to it. I did not follow the
more slender vessel beyond the posterior ovarian. Between the
left renal vein and the posterior bifurcation of the postcaval were
four stout intercostal veins, of which two lay above the entry of
the right ovarian, which enters the postcaval in common with
the third of these four veins. The lumbar parietal veins flow
into the common iliac vein on either side. The ovarian arteries
escape from the aorta just at the bifurcation of the postcaval, as
shown in text-figure 132, p. 499.
In Pseudochirus peregrinus the veins in question are much like
those of Trichosurus vulpecula, but show differences of detail.
The renals are but slightly asymmetrical. Into each renal opens
a vein at right angles, which is compounded of two running along
the ureters. These become separate ata very short distance from
the renal vein. ‘They appear to cross the spermatic vein without
forming an anastomosis with it, and they do not anastomose with
each other. The spermatic veins are symmetrically paired and
flow into a postcava some way below the renals. There are two
intercostal veins lying between the influx of the spermatics and
the left (lower) renal. On one side of the body I noted an
anterior lumbar parietal vein, which passes along the lower border
of the kidney at right angles to the anterior spermatic vein,
from which it arises close to the renal vein. The posterior lumbar
parietal veins flow perhaps rather into the iliacs than directly
into the postcaval stem, but are just at the junction of the
two.
In Onychogale lunata both anterior and posterior spermatics
were present, though I am uncertain as to an anastomosis
between them. The posterior spermatics were very asymmetrical,
the left being much above the right. The latter, indeed, joined
the postcaval only just in front of the bifurcation to form the
iliac veins. The renals were also as usual asymmetrical.
In Onychogale frenata the typical Marsupial arrangement was
met with in all its essentials. There were nevertheless slight
differences in detail between these two Marsupials. The renal
veins although asymmetrical were not very much so, the right-
* Tt is impossible to fix accurately, without embryological data, which part of
the postrenal section of the postcaval is here dealt with.
502 MR. F. E. BEDDARD ON THE [Apr. 27,
hand vein being only slightly above the left. The individual
dissected was a rather large young one, but from the pouch; but
it does not appear likely that it would have retained any feetal
characteristics. On each side the usual two veins open into the
renal. One of these, the outer renal, seems to open directly into
the kidney, as I have noticed in 7richosurus vulpecula. The two
veins do not form any anastomoses that 1 could detect, and
neither of them become connected with the posterior spermatic
veins. The latter are quite symmetrical. The left suprarenal
vein opens into the renal vein just before the latter debouches
into the postcaval.
Text-fig. 133.
Postcaval vein and its branches in Dasyurus maug@i (A) and
Onychogale frenata (B).
sr. Suprarenal vein. Other lettering as in text-fig. 132.
In another specimen of Onychogale frenata there were some
slight differences. This example was an adult female with a
small young one in the pouch. The renals were perhaps rather
1909. | POSTCAVAL VEIN IN MAMMALS. 503
more asymmetrical. Both renals, it ought to be stated, were
single veins, as in the last specimen. The suprarenal of the left
side entered the postcaval just at its junction with the left renal.
The posterior ovarian veins were nearly but not absolutely
symmetrical, the right-hand vein being just a shade above the
left. The “ieee ovarian veins were double on each side, and
certainly anastomosed with the posterior ovarian on each side,
where they passed dorsal to it. The two veins—closely following
the ureters, as is the case in other Marsupials—were connected to
each other by numerous anastomoses, and the conditions were
the same on both sides of the body. Between the entrance of
the renal vein (left) and the point of entrance of the two posterior
ovarian veins I counted three rather slender intercostal veins, of
which two were distinctly on the right side of the postcaval, and
one (the last) as distinctly on the left side. The lumbar parietal
veins opened into the iliac a considerable distance from the bifur-
eation of the postcaval, and were directed anteriorly in their
course.
In Macropus parryi the typical Marsupial conditions are met
with. The example which I dissected was in very good condition
for the examination of the veins; for these vessels were turgid
with blood and the smaller branches could be followed with ease.
The renal veins are roughly symmetrical: there were two of them
on the right side and one on the left; the left renal vein divided
into three before entering the kidney. The ovarian and its
parallel vessel, which empty themselves into the renal vein on
either side, were well developed, and each couple of vessels were
connected by more than one anastomosis. J am not quite certain
whether they were also connected with the posterior ovarian
veins flowing into the posterior part of the postcaval. In any
case the connection must be by means of minor twigs; for the
main vessels could be seen easily to pass dorsally to the posterior
ovarian trunk on each side on their way from the generative
organs. One of the two anterior ovarian veins becomes ¢ connected
by a transverse branch with the common iliac vein just before
the union of the latter with its fellow to form the postcaval.
The position of this vein, which runs transversely into the
iliac, is very suggestive of the posterior lumbar parietal vein of
other Mammals. It is, however, certain that in the present
species the vein does not bring back blood from the parietes; it
is merely a junction as already described. I could not find a
vein corresponding to this on the right side. The left suprarenal
vein had an antero-posterior course opening into the left renal.
The two posterior ovarian veins were asymmetrical, the left
joining the postcaval considerably higher up than the right. The
left vein had furthermore two branches, also connecting it with
the posteava lower down. The lowermost of these was a stout
branch which gave off a backwardly running twig to the parietes ;
it may be that this vessel is really the homologue of the lumbar
parietal above referred to. I noticed nothing of the kind in
504 MR. F. E. BEDDARD ON THE [Apr. 27,
connection with the right posterior ovarian vein. Between the
right renal and the right posterior ovarian vein four median inter-
costal veins opened into the postcava. These veins appeared to
me to be particularly slender.
In Macropus hagenbecki the renals were nearly symmetrical,
and so also were the posterior spermatics; the anterior spermatic
veins were present.
Text-fig. 134.
Le;
Postcaval vein and its branches in Macropus parryi (A) and
Macropus agilis (B).
In the latter the spermatic arteries are shown emerging from the concealed
aorta just below the posterior spermatic veins.
Lettering as in text-figs. 1382, 133.
Macropus agilis agrees very closely with I. parryi. The lett
suprarenal flows into the left renal near to its entrance into the
posteaval. The two vessels accompanying the ureter appeared to
me to anastomose, one of them at any rate and that on the left
side, with the posterior ovarian vein. The posterior ovarian veins
1909. | ; POSTCAVAL VEIN IN MAMMALS. 5O5
were symmetrical and had no secondary connections with the
posteaval*, The intercostals were very slender as in Macropus
parryy and apparently paired at their entrance into the postcaval.
Into the left common iliac opened a medianly situate caudal
vein. The ovarian arteries, contrary to what I have recorded in
Trichosurus fuliginosus, emerged along with the ovarian veins.
On the left side I counted two intercostal veins between the
renal and the posterior ovarian,
Dasyurus manger (see text-fig. 133 A, p. 502) shows some slight
variations in the postcaval branches, which do not, however, in
any way detract from the quite typically Marsupial arrangement
of the veins in that animal, I have examined most of these
veins in five examples. ‘The renal veins in all were symmetrical,
the right being considerably in advance of the left. The posterior
spermatic veins were also very nearly if not absolutely symmetrical,
The usual two veins on each side entered the renals; but in one
specimen at any rate the outermost of these veins seemed to enter
the kidney itself, there being thus a kind of suggestion of a renal
portal system. IJ have noticed this same connection with vessels
actually within the kidney in other animals, and it may possibly
be the persistence of an embryonic condition. The importance of
these anterior spermatic veins appears to vary in individuals, and
I think that they do not always anastomose with the posterior
spermatics. The left suprarenal sometimes opens into the left
renal and sometimes into the postcaval vein direct.
Among the Edentata I have chiefly examined the Dasypodide.
It is already clear from the investigations of Hochstetter that
the genus Dasypus is characterised by a postrenally divided post-
cava. ‘l'wo species figured and described by Hochstetter + (whose
figures are copied by McClure {), viz. D. setosus and D. novem-
cinctus, differ, however, in certain details. In Dasypus setosus
the postcava is double at an earlier position than in the other
species of Dasypus. Immediately after the embouchure of the
two renal veins the postcava is divided in D, setosus. The division
occurs lower down in WD. novemcinctus. Moreover, in the
former species another character of this genus is shown in a more
marked fashion, 7. ¢. the caudal vein forms a more complex rete
of vessels. Finally, while in D. setosus only the left renal vein is
double, both are double in D. novemcinctus. These peculiarities
may of course be individual. In the generalities of the dis-
position of the veins concerned I can assert that Dasypus vellerosus
resembles its congeners. There are, however, differences of detail
which are worth noting, though some of them may of course be
individual.
* It seems probable that the secondary connections described above in Macropus
parryi are explicable on the same grounds as those urged in explaining the asym-
metry in Vrichosurus fuliginosus. The “connections” are probably to be looked
upon as intercostal veins, and one of them (see text-fig. 134 A) has a branch to the
parietes.
+ Morph. Jahrb. ¢. ¢.
ft Anat. Anz. xxix. 1906, p. 376, fig. 3, p. 377, fig. 4,
Proc. Zoou. Soc,—1909, No, XX XIII, 33
506 MR. F, E. BEDDARD ON THE (Apr: 27,
In Dasypus vellerosus (text-fig, 135 B) the renal veins are
asymmetrical. The right arise in front of the left, as is so very
usual among the Mammalia. On the right side there is one
principal renal vein. This is reinforced by two others, but
Text-fig. 135,
A
Postcaval vein and its branches in Tatusia peba (A) and Dasypus vellerosus (B).
A.sp. Spermatic artery. Ca. Caudal plexus. mes. Mesenteric artery.
ov. Ovarian arteries.
Other lettering as in text-figs. 132, 133.
these latter rather slender trunks do not reach the kidney by
a direct route. They open into the larger of the two veins
which arise in common from the principal renal vein and descend
1909. ] POSTCAVAL VEIN IN MAMMALS. 507
along the ureter. Their course is thus somewhat oblique. In
this we have an exact resemblance to ). novemcinctus. On the
left side the renal vein is quite single (it is double in both of the
species already mentioned) and it gives off just before entering
the kidney two veins like those of the opposite side of the body.
The left suprarenal vein enters the postcaval, just anteriorly to
the influx of the left renal. The point at which the postcaval
bifureates is behind the renal veins and seems to me to be pretty
well intermediate in position as compared with the two forms
studied by Hochstetter. The rete formed by the candal vein is
large, as in Dasypus setosus. It is a complex rete mixed up with
the caudal artery in a way which I have not disentangled.
I have also had the opportunity of examining a fourth species
of Dasypus, viz. D. villosus. Of this species I have dissected two
examples, but am only able to say something concerning the
details of the postcaval vein in one. Of the other I can only say
that it showed the typical condition of this genus. It is of course
important even to say this. For in some mammals which show
a double postrenal postcaval (e. g. Jctonyx*) there is variation,
some examples being normally Hutherian without a postrenal
doubling. In one example of D. villosuws the veins in question
were simpler than in the other types hitherto dealt with. The
renals were asymmetrical as in the others; but there was only a
single renal vein on each side. These two veins appeared to me
to be of about the same size. The left suprarenal vein entered
the left renal only just before its connection with the postcaval.
From each renal arose a single ovarian vein descending parallel
to the longitudinal axis of the body as in the other species. On
both sides the lumbar veins were two, opening each into the
divided region of the postcaval. I did not note these veins in the
other species.
I am also able to add to what is known concerning the post-
caval vein and its branches in the Armadillos some notes upon
these veins in Lysiwrus unicinetus, a species of which I have
dissected one example which was a male. In this Armadillo the
division of the postcaval is more marked than in any species of
Dasypus (among those referred to in the present communication).
The division just extends anteriorly to the influx of the renal
veins; and the conditions which obtain in this genus are there-
fore like those which occur in Mephitis tf occasionally, where,
however, it is only one renal (the left) that opens into the divided
portion of the postcaval. In the Armadillo with which I am now
concerned both renal veins are thus separate from each other.
Another noteworthy point about this species is the greater calibre
of the renal vessels on the left side of the body as compared with
those of the right. They were vast in the specimen which I
dissected and turgid with blood. On the left side of the body
* Of, Beddard on Glalidia &e., P. Z. S. 1909, p. 491.
+ P, Z. 8. 1909, p. 492.
Baie
508 MR. F, E. BEDDARD ON THE [ Apr. 27,
there are in fact two renals, of which the lower is the larger; the
two join before entering the substance of the kidney and may be
also said to rejoin before entering the left half of the postcaval
vein. The diameter of the lower renal is greater than that of the
posteaval into which it flows. The left suprarenal vessel just
Text-fig. 136.
Postcaval vein and its branches in Centetes ecaudatus (A) and
Lysiurus unicinctus (B).
J. Junction between two postcave in Centetes.
joins the upper renal vein before the latter joinsits fellows. From
the lower renal vein two veins arise, of which the outer is the
spermatic ; it soon divides into two trunks, but just before this
1909.] POSTCAVAL VEIN IN MAMMALS, 509
sends off a branch to the inner vessel which runs along the ureter.
On the right side the renal vein was quite single. With it are
connected two veins as on the opposite side; but these as well as
the renal vein were of less calibre than their fellows on the left.
I noticed a double lumbar parietal vein on the left side; but
have no notes concerning a corresponding vein on the right. I
also observed a caudal plexus in the pelvic region, which is so
characteristic a feature of the venous system of the genus
Dasypus. In the large tract of postcaval, which is double, it is
clear that this species comes nearer to Dasypus setosus than to
the other species of that genus which have been described.
I did not find posterior spermatic veins in this Armadillo, and
am sure that I have not confused such veins with the lumbar
parietal veins already referred to.
I have already mentioned in a preliminary note* that the
venous system of the genus Vatusia is constructed upon the same
plan as that of the genus Dasypus, instancing the species Vatusia
kapplert. Of that species I have accurate notes upon the left
side of the body only. The postcava is double at some little
distance behind the influx of the renal veins and just in front of
the origin of the ovarian arteries. The left renal vein is single
and enters the postcaval behind and not in common with the supra-
renal of its side, Just before the renal vein reaches the kidney,
or, to speak more accurately, just after it has left the kidney, it
gives off a descending branch which fuses with the ovarian
vein to be described shortly and then bifurcates, one branch going
to the ovary and the other passing down the ureter. The latter
is connected with an ascending branch which I did not trace
so far forwards as the renal vein, but it evidently corresponds to
the second of two veins which debouch into the renal vein in other
Armadillos. The ovarian veins are quite symmetrical, and, as in
Dasypus novemeinctus, as figured by Hochstetter, flow into each
half of the postcava just after the bifurcation and on a level with
the emergence of the posterior mesenteric artery.
Of Tatusia peba I have more elaborate notes. The individual
which I dissected shows some differences from the specimen of
Tatusia kappleri just described. ‘The renal veins (see text-
fig. 135 A, p. 506) are asymmetrical, but the left does not become
connected with the postcaval a very long way below the right.
The renal veins are not, however, single on each side as in Tatusia
kappleri (as to right side); there are two veins on each side, of
which the more anterior is in both cases the larger. From the
lower of the two renal veins of each side depend two veins which
are again fairly symmetrical on both sides of the body. These
are shown in the figure (text-fig. 135 A), and they will be seen
to join and rejoin in a plexus-like fashion. The outer branch is
ovarian and the two inner follow the ureter. I do not think
that there are ovarian veins connected with these and with
* Amer, Journ. Anat. 1907, p. 111.
510 MR. F. E. BEDDARD ON THE [Apr. 27,
the main postcaval trunk. On one side I found a small vein which
T regard as lumbar; but I could see nothing on the other side.
On the other hand, an additional vein to the ovary was connected
with the postcavals far down and in the pelvic region close to the
ovaries. The postcaval becomes a double vein some way after
the renals and the position of the point of bifurcation seems to be
much as in Zatusia kapplerit. I have no notes as to a caudal
plexus in this Armadillo, and I do not think that I should have
failed to note it were the plexus present. At the same time I
feel unable to assert positively that it is absent.
It is clear, therefore, from what has been said and from a
comparison of the facts which I am able to bring forward here
with those noted by other observers, that the Armadillos as a
family are to be characterised by the invariable presence of a
divided posteaval, but the actual point at which the division
occurs differs. Again, in all Armadillos the sexual veins are
symmetrical and connected with the renals, while accessory veins
to the gonads are not invariably present; when present they
occur some way down the postcava and are symmetrical, The
caudal plexus is not universal, but is characteristic of the genus
Dasypus alone of those examined.
While it appears to be clear, from the investigations of Hyrtl™,
Hochstetter 7, and myself %, that the divided postcaval vein is a
clistinctive feature of the Ar madillos, occurring, without exceptions
as to species or individuals, in all of the four genera which are at
the present time known, the facts are otherwise among the
Insectivora. In both Hrinaceus algirus and EH. europeus there
is a single postcaval vein which is undivided until it divides in
the eral way to form the two iliac arteries, or there may be
(Z. ewropeus) a divided posteaval as in Centetes. On the other
hand, in the genus Centetes the postcaval vein is broadly as in the
Armadillos, though, as will be seen presently, there are differences.
Tn Erinaceus algirus—to commence with the simpler form seen in
Insectivora—the renals are asymmetrical, the left lying, as is the
rule in such cases, below the right vein. The left suprarenal vein
opens into the left renal and just above the entry into the latter
vein of the ovarian vein which runs down along the ureter. The
corresponding ovarian vein of the right side enters the postcaval
at a point about on a level with the entrance of the left renal:
there is thus a kind of symmetry in the ovarian veins such as
occurs in other animals, for instance in Lagostomus trichodactyluss,
Paradoxurus her maphr oditus |, «we. The two lumbar parietal
veins, so constant among mammals, open symmetrically into the
iliac veins just below the partition of the postcaval.
In an example of Hrinaceus europeus I observed a slight
difference from the state of affairs here recorded in Z#. algirus,
which I do not for a moment attribute to specific differences, but
* Denkschr. Ak. Wien, 1855. + Morph. Jahrb. xx. t Supra.
§ Vide p. 516 of the present communication.
|| Beddard on Galidia &c., P. Z. S. 1909, p. 489.
1909. } POSTCAVAL VEIN IN MAMMALS. 511
rather regard as a mere variation of an individual character. In
the European Hedgehog I found that the renal veins were quite
or very nearly symmetrical. It ought also to be stated that the
posteaval vein lay posteriorly to the right of the aorta, as is the
normal position in EKutherian Mammals. In a second specimen
there was a divided postcaval as in Centetes; but I am unable to
give details.
In Centetes ecaudatus (see text-fig. 136 A, p. 508) I found the
conditions characteristic of the Armadillos somewhat exaggerated.
In a specimen of that Insectivore which I dissected some two
years ago the postcaval was formed of two parallel vessels from
a point slightly in front of the influx of both renal veins anteriorly.
A larger region was therefore double that in Dasypus &e. The
renal veins were single on both sides, the complication of these
veins so often seen among the Armadillos being in Centetes quite
undeveloped. The two divisions of the posteaval after the influx
of the renal veins were separated somewhat widely as in Dasypus
&e.; but this separation was not complete. For two veins
bridged over the intervening space, as is shown in the figure
(text-fig. 136A). The first of these was slightly behind the
left renal, and each slender vein joined its fellow to form a short
backwardly running median vein. Further back still, just in
front of the iliac veins, there was another small median vein
formed by the union of two similar branches. The posterior
mesenteric artery emerges from the aorta in front of the anterior
bridge. The conditions thus shown are not seen in the Armadillos.
But they exactly correspond (in that there are two bridges between
the right and left postcavee) to a stage in the development of these
veins in Hehidna aie by Hochstetter *, whose figures have
been copied by McC ‘lure +
Such anastomoses nilso% occur in Didelphys as stated by McClure.
It is perhaps, therefore, possible to say that the presence of such
anastomoses uniting the postrenal section of the postcaval, which
is, as I presume, formed at least more anteriorly by the two sub-
cardinals of the embryo, is an archaic character; for during the
development of Didelphys such anastomoses occur of which there
are in that Marsupial, as already stated, frequent traces in the
adult in various positions and of differing degree, many of which
have been deseribed and figured by McClure in his paper to which
reference has been made so often in the present communication,
But without actual developmental facts relating to Centetes it is
of course dangerous to press any such comparisons very far. For
it is equally possible that the anastomoses in question are a
remnant of the cardinal collateral system. In Centetes, as in
Erinaceus and other primitive mammals, each renal vein received
from the hinder region of the body two veins which course along
the ureter; one of these, as in other animals is the ovarian vein.
* “Monotremen u. Marsupialier,” in Semon’s Zool. Forschungsreise in Australien,
Bd. ii. Lief. iii.
+ Amer. Journ. Anat. ii. 1903, p. 400, fig. x.
512 MR. F. E, BEDDARD ON THE (Apr, 27,
The lumbar parietal vein of one side at any rate emerged from
the parietes in a plexus form and entered the right half of the post-
caval some distance below the entrance of the renal of the opposite
side of the body and a fortiori below the entrance of the renal of
its own side of the body. The left suprarenal vein is formed of
three principal twigs which leave the suprarenal body ; it enters
the postcaval in front of the entrance of the left renal and not in
common with that vein. It is therefore to be noted that in this
animal the suprarenal and ovarian veins do not lie in the same
straight line and present the appearance of being portions of
the same original vein (the left posterior cardinal) as they do
in various Marsupials. Although I am of my own knowledge
unable to do more than state that one of the two examples of
Erinaceus europeus which I dissected showed the divided post-
caval characteristic of the Edentates, Monotremes, &c., the con-
ditions which actually obtain in such a specimen with a divided
postcaval have been figured and described by Hochstetter *.
This figure shows that the arrangement of the veins in Hrinaceus
1s In some respects different from that which I have described and
here figure in Centetes (see text-fig. 136 A, p. 508). They both
agree, however, in that the division of the postcaval extends further
forward than in the Armadillos. The postcaval is formed of two
veins in both species up to a point just anterior to the opening
into it of the right (and anterior) of the two renals. There
appears, moreover, to be a junction between the two halves of the
postcaval in Hrinaceus corresponding to the posterior of the two
transverse trunks of Centetes. Otherwise one is struck rather
by the differences than by the points of likeness which the post-
caval venous systems in these two types show to each other.
The kidneys and their veins are more symmetrical in Hrinaceus.
From the left renal arises only a single vein, which is the ovarian
of Centetes. It is unaccompanied in Hrinaceus by a second vein
coursing along the ureter. Moreover, this vein in Hrinaceus
anastomoses with a posterior ovarian vein as in the genus Dasy-
urus (im some specimens). The condition is, in fact, more
“‘ Marsupial” than is that of Centetes. On the right side there
is no symmetry in the ovarian vein. It flows into the postcaval
vein of its side, and not into the renal asin Centetes. It, however,
joins the other ovarian vein below. It is to be noted that this
example of Hrinacews, which Hochstetter speaks of as “‘abnormal”
in the disposition of the postcaval vein, agrees in this last particular
with a specimen of Hrinaceus algirus examined by myself and
reported upon in the present communication. For in the latter
example the ovarian veins show precisely the same asymmetry.
It is clear from the additional facts which I am able to bring
forward in this paper, that the postcaval venous system of the
Insectivora has not so completely rid itself of its primitive paired
character as it has in the Carnivora, where apparently only vestiges
* Morph. Jahrb. Bd. xx. 1893, Taf, xxiii, fig. 24.
1909.] POSTCAVAL VEIN IN MAMMALS. 513
remain of the double postcaval*, The Edentata, Insectivora, and
Carnivora can be arranged in a series in the order in which they
have just been named in respect of the double character of the
posteaval veins, 2. e. the more or less complete persistence of both
of the subcardinal veins or cardinal collateral veins. It is doubt-
less important to note that the Rodentia, which are equally
primitive in the azygos veins, show no traces of any likeness to
the Insectivora in respect of the postcaval.
The condition characterising the adult Centetes and Dasypodidee
is also very well seen as a temporary feature of the foetal Mole.
Ata certain stage figured by Messrs. Soulié and Bonne 7, there
are two thick veins which unite inferiorly and which I presume
represent the cardinal collaterals as described by McClure. A
small branch of the renals which is not lettered by these authors
seems to correspond to the real cardinals. Between the preserved
cardinal collaterals (in the 12°5 mm. long embryo described by
Soulié and Bonne), which are thick veins, is a medianly situated
single vein of some calibre, which may perhaps be the equivalent
of the subcardinals as described by McClure in the American
Opossum, or possibly be the equivalent of the median prolongations
of the commissural vessels in Centetes. This soon disappears as
an important vessel, but remains as a slight commissure. ‘The
authors speak of the persistent postcaval as being formed in the
postrenal region by the ‘right internal vein of mesonephros,”
which is not homologised with any particular division of cardinals.
There seems also at a certain stage to be a caudal plexus which
heightens the likeness to the adult Armadillo, The developmental
phenomena argue at least a general similarity with other Mammals
as described by other authors.
As to the Ungulata, I have examined the postcaval and its
branches in several examples of Hyrax capensis and in one of
H. dorsalis. The general arrangement in both species appears to
be as follows :—The renals are asymmetrical as usual, the right
opening into the postcaval above the left. The postcaval post-
renally lies as usual to the right of the aorta. The ovarian vein
on the left side opens into the renal, but on the right into the
postcaval in line with the orifice of the left renal. There is thus
a kind of symmetry between the two ovarian veins such as is
met with in the Rodents, some Carnivores, &e. In one specimen
(of capensis) I noted more particularly the branches of the right
ovarian vein. This vein shortly after leaving the postcaval (it
is simpler for purposes of description to disregard the direction
of the blood-flow and to treat these affluents as branches) divided
into two equisized veins. The anterior of these was directed
forwards and ran along the body-wall giving off branches to
the parietes. It ran about parallel to the postcaval and its
course suggests an anterior section of the postcardinal. The
* See Beddard, P. Z. S. 1909, p. 491, text-fig. 131.
+ “Recherches sur le développement du systéme veineux chez la taupe,” Journ.
de V Anat. et Phys. xli. 1905, p. 1.
514 MR. F. E. BEDDARD ON THE [Apr. 27,
posterior branch ran in the same straight line as the anterior
branch and divided into two principal trunks, of which the
anterior supplies the ovary itself and the hinder the oviduct.
In two other examples I found this branch just as plain. It is
remarkable that it was at least not so obvious on the left side ;
indeed I made no notes at all as to its occurrence on that side of
the body. It is possible that the restriction to, or at least the
more marked development upon, the right side of the body is con-
nected with the persistence of the right azygos vein only (as a
rule) in this Subungulate.
McClure has discovered * the interesting fact that in Zragulus
meminna the posteaval is divided postrenally as in Armadillos &e.,
and I have been able to confirm f him by the examination of three
individuals. Since the publication of my note referred to I
haye examined two other examples of Zragulus meminna which
presented an identical disposition of the vena cava posterior.
Furthermore, I have still more recently dissected an example of
Tragulus stanleyanus in which the same bifurcation of the post-
caval occurred, and in the same way the genital veins arose sym-
metrically from the region of the postcaval.
The Lemuroidea.—In the course of some notes upon the
anatomy of Chiromys madagascariensis = I have pointed out that
within the group of Lemurs considerable variations exist in the
tributaries of the postcaval vein. It seems to me to be evident
—though I did not point it out in the memoir to which I have
referred—that Microcebus has retained the Marsupial and Kdentate
character in its venous system, in that the spermatic veins arose
more or less symmetrically some way down the postcaval and that
both were connected with the renal veins by a slender vessel on
each side, which seems to me to represent the anterior spermatic
vein of Marsupials, but in this Lemur (icrocebus) mm the course
of disappearance. On the other hand, in Chiromys (of which
Lemur I reported upon two examples in the paper referred to and
have here to report upon a third $) only the posterior spermatic
veins in the male at any rate are left.
A study of the posteaval and its branches in Lemur catia shows
that this region of the venous system is like that of MZicrocebus.
The two renals are single on both sides, but asymmetrical in the
usual way. Into each renal flows on the posterior side an anterior
parietal vessel, which corresponds, as I believe, to one of the two
veins on each side which accompany the ureter in the Marsupials.
On the left side there was also a small vein like that in crocebus
closely accompanying the posteaval and flowing into it above close
to its junction with the left renal. Posteriorly, I think, but am
not quite certain, it joins the spermatic of its side. In this case
the likeness to Mierocebus is quite close. The two principal and
* “The Postcava of an Adult Indian Chevrotain,” Anat. Anz. xxix. 1906, p. 375.
+ Amer. Journ. Anat. vol. vii. 1907, p. 111.
+ P.Z.S. 1908, p. 694.
§ Vide infra, p. 521.
1909. | POSTCAVAL VEIN IN MAMMALS, 515
posterior spermatic veins flow into the postcava quite far back
near to the bifurcation of that vein. They are quite symmetrical.
Between the influx of the left renal above and of the spermatic
veins below I counted four unpaired intercostal trunks. The
iliac veins were asymmetrical, as is shown in the drawing (text-
fig. 137), the internal arising as one trunk from the left external
iliac.
Text-fig. 137.
Postcaval vein and its branches in Microcebus smithii (A) and Lemur catta (B).
Ao. Aorta. Other lettering as in text-figs. 132, 133.
In so far as concerns the Rodentia, the Rabbit is naturally very
well known * ; and Hochstetter has furthermore called attention T
to several variations in the venous system as affecting the tribu-
taries of the postcaval. On the whole, it would appear that the
* Krause, ‘ Anatomie des Kaninchens,’ Leipsic, 1884; Haswell & Parker, * Text-
book of Zoology,’ London; aud many similar works.
+ Morph. Jahrb. Bd. xx. pp. 585, 586, figs. 7, 8, 9.
516 _ MR. F, E, BEDDARD ON THE [Apr. 27,
several branches of the postcava in that animal are as in the Hys-
triciform Rodents, of which I give a certain number of details
presently. In two out of the three venous systems figured by
Hochstetter the left spermatic vein entered the left renal vein,
while the right spermatic vein entered the postcaval a good way
below the entrance into it of the two renals. A very remarkable
state of affairs was shown in a third example which I have not
been able to parallel in any Rodent dissected by myself. In this
animal the left spermatic artery entered the left parietal lumbar
vein.
Of Hystrix cristata 1 have examined the postcaval vein and its
branches in three examples, in all of which the conditions were
much the same but not precisely identical. All the renal veins
showed the usual asymmetry, which need not be further particu-
larised. In all, the left ovarian vein entered the left renal vein ;
in two of the individuals the right ovarian vein entered the post-
caval a good way below the entrance of the left renal; in the
remaining specimen the point of entrance was higher up on a level
with the entrance of the left renal, as in the Myomorpha generally.
In one specimen (I have no notes upon the other two as concerns
this vein) 1 observed the right lumbar parietal vein to enter the
corresponding iliac vein and not the main trunk of the postcaval.
In another specimen (again I have no notes upon the matter in
the remaining two) the caudal vein was asymmetrical and entered
the left iliac.
Chinchilla lanigera is quite like Hystrix cristata; in both of
two specimens the renals were asymmetrical and the left genital
vein entered the left renal, while the right genital vein entered
the postcava some way down. An example of Dasyprocta cristata
showed the same features. A specimen of the Capybara, Hydro-
cherus capybara, was identical, save for the fact that the renal
veins were fairly symmetrical. The left suprarenal vein, it should
be mentioned, opened directly into the postcaval vein above the
renals. In a second specimen of this Rodent the renal veins
were also symmetrical. A specimen of Aulacodus swindernianus
showed also symmetrical renal veins ; the ovarian veins were as in
the last three genera. Of the Viscacha, Lagostomus trichodactylus,
I have dissected the postcava in two specimens. In one the
renal veins were about symmetrical; in the other the right was a
little in advance of the left. In both specimens the ovarian vein
of the left side opened into the corresponding renal, and of the
other side into the posteava opposite to the left renal.
The posteaval and its branches in Hydromys chrysogaster (text-
fig. 138 B) are as follows:—The postcaval itself occupies the usual
position to the right of the aorta. The renals are asymmetrical,
the left vein opening below the right in position. The left
suprarenal vein receives, after leaving the suprarenal body, three
parietal veins, two anteriorly and one posteriorly. The left
ovarian vein enters the left renal, and the right ovarian vein
enters the main trunk of the postcaval not a very great way
1909. ] POSTCAVAL VEIN IN MAMMALS, 517
behind the entrance of the left renal, the prevailing Kutherian
arrangement thus occurring. The lumbar veins are asymmetrical,
the left vein debouching into the postcaval above the right.
Text-fig. 138.
Postcaval vein and its branches in Sciwrus prevosti (A), Hydromys chrysogaster (B),
and Dipus hirtipes (C).
Lettering as in text-figs. 132, 133, 137.
Gerbillus egyptius and Zapus hudsonianus (text-fig. 139, A, B,
p-519), whichare both members of the group Myomorpha, but which
are placed by Mr. Thomas * in quite different sections of that group,
show an interesting point of similarity, of which possibly it is easy
to exaggerate the importance as a mark of similarity. Posteriorly
in both animals the postcaval vein lies accurately above the aorta
* P. Z.S. 1896, p. 1016.
518 MR. F, E, BEDDARD ON THE [ Apr. 27,
(2. e. below” when the animal is dissected, as is usual, from the
ventral surface) for some distance in front of the point where it
emerges from between the divergent iliac arteries accurately in
the middle line. This is, of course, only an exaggeration of the
prevailing EKutherian condition. Still it is notewor tihy that a new
kind of symmetry in the relation of vein and artery precisely the
reverse of that characteristic of the Marsupials should be visible in
these Murines, Finally, in both of these species the spermatic
vein entered the iliac vein of its side, and was thus not directly
connected with the postcaval or renal, as is the usual state of
affairs in Kutherian Mammals.
In Georhychus capensis (text-fig. 139 C) the veins with which I
am concerned in the present communication are different from
those of some other Rodents. The renal veins are not situated
opposite to each other, the left being below the right. The right
ovarian vein is symmetrical with the left renal and passes, like it,
almost at right angles to the postcava, the ovary of that side
being situated to the outside of and not much behind the kidney.
It gives off two branches which run at right angles to it and
parallel with the postcava along the ureter and the oviduct. On
the left side the ovarian vein joins the left renal, but before it
joins receives a vessel running along the oviduct. The renal
vein nearer to its embouchure into the postcaval receives a slender
vein which runs along the ureter. Posteriorly there is a caudal
vein which opens into the left of the two iliac veins which together
form the posteaval. Dipus hirtipes (text-fig. 138 C, p. 517) does not
differ much from Georhychus. But the right ovarian vein flows
into the postcaval rather nearer to the right renal.
Graphiurus murinus is, i many respects, like Georhychus
capensis. The renals are even more asymmetrical, the space of
postcava left between their respective mouths being greater. The
left suprarenal vein enters the postcava opposite to the right
renal. Between the two renals enters the right ovarian vein ;
this is, as in Georhychus, at right angles or nearly so to the post-
caval stem, and, as in the Rodent mentioned, receives an ascending
vein which runs along the oviduct. On the left side of the body
the ovarian vein enters the left renal; but the latter vein, instead
of being at right angles with the posteava or sloping poster iorly,
1S directed rather ‘anterior ly. This ovarian vein receives an
ascending branch as does the right ovarian vein.
In Sciurus prevosti (text-fig. 138 A) the renal veins were
asymmetrical in the usual fashion. The ovarian veins were
symmetrical and situated far back, flowing into the postcaval not
far in front of the union of the iliacs. Between these veins and
the left renal there were three unpaired intercostal veins. Sciwrus
cinereus was the same, except that the two spermatic veins
appeared to open into the postcava by a common stem. In front
of this were two lumbar parietal veins, of which the left-hand
one was rather the anterior. In Sciwrws maximus the spermatic
veins had the same symmetry and posterior origin.
1909. | POSTCAVAL VEIN IN MAMMALS. 519
The posteriorly situated point of opening of the spermatic and
ovarian veins in the genus Sciurus is, so to speak, a less exaggerated
condition of these veins than that found in Gerbillus and Zapus,
and is suggestive of the conditionswhich appears to be universal
among the Marsupials.
In Notiomys cervinus (of which I have dissected two examples,
both females) the renal veins show the usual asymmetry, the
Text-fig. 139.
Postcaval vein and its branches in Zapus hudsonianus (A), Gerbillus egyptius (B),
and Georhychus capensis (C).
Lettering as in text-figs. 132, 133, 137.
right entering the postcava higher up than the left. In both
specimens the right ovarian vein entered the postcaval at a point
pretty well opposite to the entrance of the left renal. The left
520 MR. F. E. BEDDARD ON THE [ Apr. 27,
ovarian vein, on the other hand, entered the left renal. In one
of the two examples which I dissected, and of which I took more
elaborate notes, a small vessel running along the ureter also entered
the left renal. In this specimen the left suprarenal vein before
entering the left renal received a branch from the parietes ante-
riorly, and there were two large lumbar parietal veins entering
the postcava before its posterior bifurcation and asymmetrically ;
the left vein entered the postcava higher up than the right.
Although these vessels ran towards the uterus, they did not,
as it appeared to me, receive any affluents from any part of the
generative system.
In Mus coucha the conditions which obtain are very much lke
those seen in Votiomys. The two renals are asymmetrical in the
same way and the ovarian veins have precisely the same connec-
tions, which, on account of their identity, I need not recapitulate.
There are two other veins which correspond in their distfibution
to the lumbar parietal veins of Votiomys, but which are differently
connected with the postcaval. On the left side of the body the
vein in question enters the left renal to the inside of the point of
entrance of the left ovarian vein, but quite independently of it.
On the right side of the body this parietal vein has quite a different
connection, for it joins the right ovarian vein before the latter
debouches into the postcaval vein opposite to the entrance of the
left renal.
Otomys irroratus agrees with Notiomys cervinus in the dis-
position of the postcaval and its branches. The two renals are
asymmetrical, the left, as usual, being below the right. Into the
left renal flows a vein which is composed of two chief affluents, a
vein from the ovary itself and a longer vein from the uterus. On
the right side the same two veins are present, and form a common
trunk which flows into the postcava just opposite to the entrance
of the left renal. The two suprarenal veins enter the postcaval
independently of the renals. The left suprarenal has a curious
course; it actually pierces the liver on its way to join the post-
caval. Below the ovarian veins a large lumbar parietal vein on
each side joins the postcaval. These are nearly if not quite
symmetrical.
In Arvicanthis pumilus there are no great differences. The
renals are not symmetrical ; the left is joined by the ovarian vein,
while the right ovarian vein enters the postcaval a little way
below the entrance of the left renal.
The few Rodents upon which I am able to report in the present
communication very clearly show that this group is by no means
so uniform as the American Edentates or Marsupials. It is
furthermore plain that the Hystricomorpha present uniform
characters in the posteaval vein and its branches. There are no
differences of importance among the six genera belonging to this
group which I have examined.
The Myomorpha, on the other hand, are by no means so
+
uniform. Hydromys is like the Hystricomorpha ; Mus, Votiomys,
LIDS POSTCAVAL VEIN IN MAMMALS. 52)
Georychus, and some other “genera agree closely, while Zapus and
Gerbillus seem to approximate to the tee I Opinions as to
the classification of Rodents vary greatly *.
§ Some Variations of the Postcaval Trunk.
Under this heading should no doubt strictly be included a very
large proportion of the facts with which I deal in the present
communication. I limit myself, however, in this section to two
cases of apparently abnormal position of the postcava; “abnormal”
because they are not like any condition of that vein which is
characteristic of a given group or even minor subdivision of any
mammalian order. From this point of view, therefore, I cannot
include the occasional duplication of the postcaval in Hrinaceus,
since it would appear to be fairly common in that genus, and is
also a marked character of the Edentata, &c. Moreover, it might
be urged that the single postcaval lying to the right of ‘the aorta
is the abnormal condition i in Hrinaceus ewr opus.
In an example of Chiromys madagascariensis dissected in
September of last year I found that postrenally the posteaval
vein lay to the left instead of to the right of the aorta. The
normal condition of this vein has been already and quite recently
described by myself 7. “The arrangement of the spermatic or
ovarian veins as the case may be appeared to me to differ in the
two sexes, and did undoubtedly differ in the two individuals upon
which I reported to the Society. Im both individuals the right
spermatic or ovarian vein flowed into the postcava rather distally
but still some way in front of the posterior bifurcation of that
vein. In the female the left ovarian flowed into the left renal,
while in the male the corresponding spermatic vein flowed into
the postcaval a little way anterior to the entrance of the right
spermatic vein. The relative position of these veins in the
male is important to be borne in mind. In the abnormal
example of Chiromys to which I refer in the present communi-
cation, the relative position of the two spermatic veins was—in
correspondence, as I imagine, with the sinistral posteava—
entirely reversed. The right spermatic vein entered the postcava
anteriorly to the left.
There was, however, no reversion in the position of the renal
veins ; they lay exactly as in the two normal examples described
by myself. It may be worth while mentioning that in this
abnormally veined Chiromys the small anterior lumbar parietal
vein, to which I directed attention as occupying in the male the
position of the ovarian vein in the female, is present; it lies,
however, on the opposite side of the renal and therefore ends
anteriorly ; it is not to be confused with a suprarenal vein. This
variation in Chiromys madagascariensis seems to be exactly
paralleled in a variation of Homo sapiens referred to by
* Of. e.g, Thomas, P. Z.S. 1896, and Tullberg, Nov. Act. Upsala, 1899.
+ P.Z.S. 1908, p. 700.
Proc. Zoot. Soc.—1909, No. XXXIV.
(Su)
re
522 MR, F, E. BEDDARD ON THE [Apr. 27,
Hochstetter *, where a prevailing left posterior vena cava was
accompanied by aright spermatic vein opening into the right
renal vein and a left spermatic opening into the postcava itself.
This is the exact reverse of the normal conditions obtaining in
Man, where, as in so many animals, it is the left genital vein
which pours its blood into the left renal while the right-hand
vein enters the postcava directly.
T have only examined one example of the American Monkey
Ohrysothri sciureus, and it is rather remarkable to find that that
one specimen, not in any way selected, should show a variation ;
for so I must in the present state of our knowledge interpret the
conditions shown by the postrenal section of the postcaval vein.
The vein was in fact, as in Chiromys just dealt with, developed
on the left side instead of the right. It lay very distinctly to the
left of the aorta. Corresponding with this abnormality in the
position of the postcaval, its principal branches showed an entirely
similar position to those of Chiromys and the human abnormality
which I have compared with it. For of the two ovarian veins
the left entered the main trunk of the postcaval some little way
down the latter, while the right ovarian vein poured its contents
in the posteaval just below the entrance of the right renal vein.
T cannot but think that in the normal arrangement of the veins
of this animal the postcaval was on the right side of the aorta
and the position of the two ovarian veins was reversed, the left
being anterior to the right.
S$ Résumé and General Considerations.
It is, in the first place, clear from what has been detailed in the
foregoing pages, and from my previous papers in which the post-
caval system is dealt with, that the postcaval system of veins in
Mammals is subject to variations of an individual character.
This conclusion is not of course deduced from my own work only,
but from what has been written by others which I am able to
confirm for other animals and from additional examples of species
already known. Even in the very few individuals of certain
species which I have been able to dissect, variation has always been
noticeable. Forinstance, in three specimens of Chiromys varia-
tions occur; in four examples of J/ctonyx capensis, in three of
Hystrix cristata, in two specimens of Hrinaceus ewropeus, in two
of Suricata tetradactyla, in three examples of Dasywrus maculatus.
And these variations together nearly cover the range of variation
which the posteaval and its branches show among the Mammalia.
In spite, however, of these variations from individual to indi-
vidual of a given species, there are certain characteristics of
groups which may be laid down with some confidence, and with
more confidence—I venture to assert—from a consideration of
the new facts which I bring forward here.
* Loc. cit. p. 638. This paper contains a good many references to the literature
of venous abnormalities in man, as well as descriptions of instances.
1909. ] POSI'CAVAL VEIN IN MAMMALS. 523
Thus the Marsupials possess a general character of the post-
caval and its branches which is shared by no other group of
mammals. Furthermore, it is very noteworthy that within this
particular group the variations in important matters is but slight.
Thus the only features which differ at all from genus to genus or
from species to species are the symmetry or asymmetry of the
renals, the exact origin of the suprarenal vein of the left side,
whether from the left renal or from an adjacent part of the post-
caval, and the freedom or connection by anastomoses of the
anterior pair of veins which leave the renals on each side together,
and finally their connection or perhaps non-connection by anas-
tomoses with the posterior spermatic veins where they cross
them dorsally on their way to the testes or ovaries. It will have
been observed that it is quite impossible to distinguish the
Carnivorous from the Diprotodont Marsupials by their veins;
the whele group of the Marsupials forms an assemblage of which
the various members are closely connected in these characters.
So far as facts already known enable us to say, it seems clear
that of the Marsupials only an individual of Vrichosurus vulpecula
and the genus Didelphys, and in the latter only occasionally and
by no means typically, have preserved the earlier condition of
the mammalian postcaval system as seen in Ornithorhynchus and
in Lehidna, where the postrenal part of that vein is persistently
double, being developed out of both collateral veins of the
embryo*. ‘This condition is, however, fully characteristic of the
Edentata, which form an assemblage apparently quite as distinct
in these characters as are the Marsupials. The present com-
munication to the Society allows me to confirm the data of
Hochstetter and Hyrtl by fresh examples of species of Dasypodide.
I may also mention that in two examples of Myrmecophaga jubata
which I have recently dissected the postcaval vein was double
postrenally, though I have not made notes of sufficient elaborate-
ness to permit of a detailed account of the facts in the earlier
part of this paper. Hochstetter has also stated that MWyrmeco-
phaga and both of the Sloths, viz. Cholwpus didactylus and
Bradypus tridactylus, possess the same type of postcaval system,
and also the Old World Edentate, Janis t. Judged by the con-
ditions which obtain in the Monotremata, the Edentata are more
primitive than are the Marsupials. And, moreover, no Edentate,
so far as is known at present, shows the typical Eutherian con-
dition of the veins in question. It is a very singular fact that
among the Marsupials the typical Eutherian condition of the
postcaval is seen in only one forin (Petaurus tanguanoides) §, just
as the Hutherian placenta is also seen in only one form. We
know so little about the Insectivora as regards their venous
* Morph. Jahrb. xx. p. 362, pl. xv., and in Amer. Journ. Anat. ii. pl. iv. figs. 17,
21, &c.
+ Morph. Jahrb. xxv. p, 621.
{ Hyrtl, Denkschr. Ak. Wien, vi. 1854.
§ Morph. Jahrb. xx. pl. xxiii. fig. 26. And, according to McClure, in one
example of Phalanger wrsinus. 'This of course may be an abnormality.
34*
524 MR. F. E. BEDDARD ON THE [Apr. 27,
system that it is rather premature at present to attempt to lay
down any statement of general application. But such facts as
we have distinctly prove that the Insectivora occupy a somewhat
midway position. They are at least commencing to lose the
double postrenal caval. Still it is very frequent. Whether
Hochstetter’s discovery of a similarly divided postcaval in
Pteropus * will prove this condition to be characteristic of the
Bats—or at least the Fruit-Bats—remains to be seen, as also
whether the Cetacea are largely or mostly thus to be charac-
terised. It can be safely said that the Carnivora yt only show
remnants of this state of affairs; but such rudiments are not
without significance inasmuch as they occur among the Arctoidea,
which are perhaps to be regarded as the most primitive among
the Carnivora. McClure’s interesting discovery, confirmed by
myself, of the condition of the postcaval in the Ungulate 7’ragulus
adds one Ungulate genus to this same category ; but as this group
has been but little studied from the present point of view, it
would be premature to base any arguments upon the facts. In
contradistinction to the groups of mammals already considered, we
have the Rodentia, Lemuroidea, and Primates, which invariably
show a postcaval vein lying to the right of the aorta postrenally
and possessing no fellow upon the opposite side. That variations
have been described does not alter the fact that these Kutherian
mammals differ from the Eutherian groups already dealt with.
It is true that here again the actual facts known require much
increase before any generalisations can certainly be made. But
the evidence is already sufficient to be seriously considered. It
will be noted that those orders of mammals which show a
retention of the Monotreme condition are admittedly primitive ;
and it may be further said generally that the greater or less
persistence of this condition is in relation to their position in
the series. Thus the Marsupials, Edentates, and Insectivora are,
in the opinion of all zoologists, ancient groups. On the other
hand, the Ungulates and Carnivora are more advanced in many
ways, while the Primates are still more specialized in some
respects. It is also noteworthy that of those types among the
more modern and specialized groups which show the character
under discussion, it is clear that they are more primitive types
than are some of their allies. The Chevrotains stand below the
Deer and Antelopes, and the Arctoid Carnivores? are nearer to
the common stem than are the Cats.
The differences observable among the genital veins are not a
* Tbid. p. 619 and references there quoted.
+ Beddard, P. Z.S. 1909, p. 491, text-fig. 13.
+ I take this opportunity of stating that in an example of Mellivora signata (in
fact, the type specimen described by Pocock, swpra, p. 394) the postcaval was divided.
The division ran up to just behind the entry of the left renal vein, as in an example
of Ictonya recently described by myself (supra, p. 491). The right renal vein entered
higher up. The ovarian veins were symmetrical and entered the divided postcaval ;
below them entered the also symmetrical lumbar parietal veins. No ovarian branches
entered either renal vein. I have figured these veins in the memoir referred to
(v. supra, p. 491, text-fig. 131 A).
1909.] POSTCAVAL VEIN IN MAMMALS. 525
little remarkable. The Marsupials seem to be alone among the
non-Monotreme Mammals in possessing invariably two pairs of
spermatic (or ovarian—for the sex of the individual does not affect
these veins) veins, one pair flowing into the posteaval direct and
the other pair, often connected on its way with the former, open-
ing into the renal veins. At least, it may safely be said that this
is Sane the prevalent arrangement. And, furthermore, this
group is also to be remarked upon as characterised by the symmetry
of the posterior spermatic veins, though here the rule is not so
universal. In other animals there would appear to be a persistence
of either one pair of these veins or of the other; while some
instances could almost seem to prove the retention of the left
posterior spermatic vein and of the right anterior spermatic vein.
The Dasypodide are apparently a group in which the posterior
spermatic veins are beginning to vanish. Both pairs occur in
Tatusia kapplert and only the anterior pair in some other forms.
The details I have already given and need not recapitulate here.
The Insectivora are a group which are in the same condition as
the Edentata. In Centetes, for example, there is no posterior
spermatic vein, while there is such a pair of veins in Hrinaceus
europeus (at least occasionally). The higher Kutherian mammals,
so far as they are known, never appear to possess both pairs of
spermatic veins. More usually, as I hope to show shortly, the
anterior spermatic vein persists on the left side and the posterior
on the right. There are, however, a few forms in which it is the
posterior pair only which remain, there being apparently in those
forms no trace of the anterior pair. Thus in 7ragulus and Sciurus
it seems to be obviously the posterior pair which are the only
spermatic veins. Furthermore, in those Carnivora (at any rate
Letonyx and Meltivora) which occasionally present us with a
divided posteaval vein, the posterior spermatic veins persist on
both sides, are symmetrical, and are the only spermatic veins. But
in examples of Letonyx where the postcaval is single and in other
Carnivora where no such duplication has been recorded, the
conditions are what may be called the typical Eutherian
conditions, 7.¢., the asymmetrical retention of veins.
This asymmetrical retention of veins seems to be connected with
the formation of the renal and postrenal sections of the postcaval
vein. As seems to be now fairly certain from the investigations
of Hochstetter *, Lewis t, McClure {, and of Soulié and Bonne §,
already referred to, that region of the postcaval in the higher
mammals is developed from the right-hand subcardinal (or
perhaps collateral cardinal) and right postcardinal only, the left
disappearing. The spermatic veins, therefore, on the left side
obviously lose their connection with the postcaval through the
disappearance of the intermediate veins. When the development
is symmetrical, as in Z’ragulus, Mellivora, &., the genital veins
* Toe. cit. pe
+ “The Development of the Vena Cava Inferior,’ Amer. Journ. Anat. i. 1902,
p. 229. t Loe. cit. § Loc. cit.
526 ON THE POSTCAVAL VEIN IN MAMMALS. [Apr. 27,
are also necessarily symmetrical. Here are both embryonic veins
preserved with their branches, naturally equivalent on both
sides of the body. But in the other cases, the spermatic vein
of the left side only retains its connection with the cardinal
(doubtless through the rich anastomoses which occur in the
embryo), and follows the course of that vein anteriorly, opening
into the renal in the left side. This point of view is confirmed
by a consideration of those cases (of which I have described two
in the present communication) where the left subcardinal or
cardinal collateral, as the case may be, is converted into the
postrenal section of the postcaval ; here the conditions of the
spermatic veins are precisely reversed, as would be expected on
this view. There remain, however, some instances which can
perhaps be disposed of on this hypothesis. There is, for example,
the single postcaval vein in the genera Seiurus and Zapus. As
to the latter, however, it is quite possible. that its median position
is significant. It may be that the two veins of the embryo which
are connected with the formation of this part of the postcaval
have both persisted and fused together, which would at once
account for the median position and for the retention of both
posterior spermatic veins. In the Squirrel, on the other hand,
the postcaval postrenally 1 is distinctly developed to the right af
the aorta as in most Mammals. Still we know nothing of the
development of the veins in this mammal, and a study of the
development might show that there was after all a fusion of right
and left veins of the embryo. Besides these cases we have the
peculiarly constant mode of connection of the spermatic veins
with the postcaval in the Murine Rodents—in many of them
at any rate, in Votiomys to take a definite example. Here we
have a symmetry between the spermatic veins but of a different
kind to that which occurs in other animals, save for a few
exceptions. The left spermatic opens into the left renal, while
the right spermatic is symmetrical with it and therefore does not
open into the renal of its side but into the postcaval opposite to
the point of entrance of the left renal. In these cases it appears
to me that we may have the disappearance of both posterior
genital veins and the retention of both anterior veins. This is, of
course, in the absence of embryological data sheer surmise; but
we already know of cases where this appears more definitely to
be the case, such as a number of Armadillos and the Insectivore
Centetes. I believe that the cases which have now been considered
exhaust the variations known to exist In mammals. From an
evolutionary point of view it seems likely that the existence of
two pairs of spermatic veins is the more primitive state of affairs,
since these veins are more numerous in the lower Amniota, and
that the prevalent arrangement of these veins as one pair is the
later state. This also (it will be observed) agrees with the mutual
relations of the groups of mammals which are here dealt with.
i
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TMG Sich COC SZ ci
USOT OSTEOLOGY OF ARACHNOTHERA MAGNA . 527
4, On the Comparative Osteology of the Passerine Bird
Arachnothera magna. By R. W. Suurenpt, M.D.,
C.M.Z.S.
[Received March 30, 1909. ]
(Plate LX VIIL*)
A number of years ago Mr. F, E. Beddard kindly sent me for
examination, from the Society’s Collections, some twelve or
thirteen alcoholic specimens of birds representing a variety of
genera and species from several parts of the world.
Tt was only very lately that I could give this valuable material
the attention it deserved, and upon comparing it with the list
submitted the following forms were found to be at my
disposal :—
No. No.
289. Cyanerpes cyanea. 306. Cereba chloropyga.
345. Arachnothera longirostris. | 718. Acanthorhynchus sp. ?
318. Jr magna. 365. Prosthemadera nove-
AQT. Leptocoma gray. hollandic«.
340. Cinnyris chalybeus 702. Entomyza cyanotis.
426. Diglossa baritula. 725. Acanthogenys rufigularis.
500. Anthreptes malaccensis. 712. Climacteris scandens.
All these specimens had been in strong spirit for many years;
some of them for perhaps twenty years, or even longer. ‘This
treatment had very much hardened all the soft tissues and
muscles, and, as some of the species are very small and delicate,
the labour of cleaning the skeletons was considerable, as 1 know
from the fact that I performed the entire task myself. In some
instances, in too many unfortunately, the extremely minute and
fragile bones—in such a species, for example, as Cinnyris chalybeus
—simply refused to hold together. Some of the specimens had
been rather roughly eviscerated, thus injuring the skeleton in
the neighbourhood of the incision. One or two were headless,
and in the case of all some enterprising and enthusiastic
ornithotomist had cut down to examine the xiphoidal extremity
of the sternum, a perfectly justifiable operation, by the way, and
doubtless to ascertain whether it was ‘‘ notched” or otherwise,
that is, passerine or trochilidine. Aside from these various
mishaps, which are comparatively few when one comes to think
how long all these specimens had been in the hands of science,
this material as prepared offers not a little worthy of study and
comparison.
The list has been kindly looked over by Dr, Chas. W. Richmond,
* For explanation of the Plate see p. 544.
528 DR. R.W. SHUFELDT ON THE COMPARATIVE OSTEOLOGY [ Apr. 27,
Assistant Curator of the Division of Birds of the U.S. National
Museum, to whom my thanks are extended for having pointed
out the changes made in the nomenclature of three of the families,
and for having brought up to date the present known habitats of
the species represented.
In this list the genera Cyanerpes, Diglossa, and Cereba belong
to the family Coerebide ; Arachnothera, Leptocoma, Cinnyris,
and Anthreptes to the Nectariniide; Acanthorhynchus, Prosthe-
madera, HLntoma Yrlt, and Acanthogenys to the Meliphagide; and,
finally, Cli imacteris to the family Certhiide.
Again, and designating the species in the list by their numbers,
we are to observe that No. 289 occurs in 8. Mexico to 8.E. Brazil
and Cuba; No. 345in the Indian Peninsula to Burmese Provinces
and Malay Peninsula; No. 318 in the Himalayas to Burmese
Provinces; No. 497 in the Celebes; No. 340 in 8. Africa;
No. 426 in S. Mexico and Guatemala; No. 500 in Siam to the
Malay Peninsula, Java, Sumatra, Bali, Borneo, and Sulu and
Tawi Tawi groups, P.I.; No. 306 in Cayenne to S.E. Brazil and
Bolivia; No. 718 in Australia and Tasmania; No. 365 in New
Zealand ; Nos. 702 and:712 in KE. and 8. Australia: and No. 725
in 8.8.E. and West Australia. Thus it will be seen that the
species are found to range through Southern Mexico, Bolivia,
South-eastern Brazil, West Indies, South Africa, widely over
India and the East Indies, Australia, and New Zealand. They
are all tropical or subtropical species, and none of them occur in
North America north of Southern Mexico; none in any part of
Europe or in the larger part of Asia.
My reasons for selecting from the list one of the species of
Arachnothera as the principal form of which the osteological
characters will be given in preference to any of the others are
that, practically, the genus is from the centre of the region of the
world’s avifauna where the other families represented find their
habitats, the regions considered being large continental areas of
the earth’s surface. Secondly, two species of Arachnothera occur
in the list, and the skeleton afforded by the specimen of 4. magna
is the most perfect of any of those obtained. Finally, in any
comparative anatomical work it is always best to select some
special form, be what it may, wherewith to compare the struc-
tural characters presented on the part of all its supposed-to-be
congeners, material for which may be at hand.
THE SKULL.— Viewing this part of the skeleton upon its superior
aspect, it is to be observed that the cranial portion is globular in
form, smooth, and is marked, mesially, by a broad shallow furrow
that runs forward and to the right, to be lost at the cranio-facial
line. This furrow is far better marked in Arachnothera longi-
rostris, and in both species affords lodgment in life for the thyro-
hyals or “ greater cornua ” of the hyoid arches. A similar groove
is found in the same locality on the skulls in the Trochilide, only
1909. | OF THE PASSERINE BIRD ARACHNOTHERA MAGNA. 529
in those birds it is better marked as a rule, and carried further
backward, having the supraoccipital prominence standing between
the furrows after the common one separates posteriorly. This
character is also a feature of the skull in Acanthorhynchus among
the Meliphagide and probably other honey-sucking species which
possess tongues, the thyro-hyals of which curl over on top
of the skull and are extensible. Cinnyris chalybeus is another
example.
In A. magna the frontal region is rather broad between the
peripheries of the orbits, and still broader in front of the
lacrymals, where the cranio-facial line is quite distinct. The
superior mandible seen from above is smooth and the culmen
rounded. This part of the skull can best be studied on side view.
Here it will be noted tbat it is gently decurved for its entire
length, which is just double of that of the rest of the skull. It
tapers very gradually to the sharp apex, while its tomia possess
clean cutting-edges. The rather large elliptical external narial
apertures open far back just beyond the cranio-facial hinge, or
rather line (see Plate), and they have no true bony partition
separating them mesially. This is entirely different from what
we find in the Humming-birds, where these mandibular narial
openings are long and slit-like. They are very large, and occupy
a mid-position on the bill in such species as Prosthemadera nove-
hollandie, Acanthogenys rufigularis, and other Meliphagide, forms
with shorter and stouter mandibles.
We find that A. magna has a capacious orbital cavity, with its
osseous walls fairly entire. The pars plana is large and thick
and faintly shows above its union with the lacrymal. Its outer
margin, forming a part of the periphery of the orbit, is, like the
rest of this margin round to the postfrontal, sharp and defined.
On its orbital side the pars plana is markedly concave, but
convex in front, while below it meets the anterior end of the
quadrato-jugal bar. The latter is almost of hair-like proportions,
very delicate, and straight. This is also the case in other species
of the Nectariniide, some few Meliphagide, and in the Trochilide.
Nearly all of these birds have a vacuity of a greater or less size
in the interorbital septum, and the openings for nerves on the
anterior wall of the brain-case, within the orbit, as those for the
first pair, are large, and in A. longirostris merge with the foramen
in the interorbital septum. The optic foramen, however, is
generally distinct, and in such a species as Climacteris scandens,
and probably its near allies, there exists no deficiency in the
orbital septum, while the brain-case above exhibits a very large
opening into the orbit.
Owing to the extreme slenderness of the osseous structures at
the roof of the mouth posteriorly, the floor of the orbital cavity
is distinctly deficient in bone, and this is the case with all the
species in this genus, as well as in some of the related forms. On
the lateral aspect of the cranium we find the postfrontal process
530 DR. R. W. SHUFELDT ON THE COMPARATIVE OSTEOLOGY [ Apr. 27
so feebly developed as to be barely noticeable, while the sqgwamosal
process below it is rather long, though extremely slender. The
valley between them is only fairly well-marked. Humming-
birds exhibit similar characters with respect to these two apo-
physes, but, taken as a whole, there is nothing, beyond the big
pars plana in one of this family, to remind us of the orbital
cavity as a whole in drachnothera. To be sure, the anterior wall
of the brain-case is deficient in bone in both, markedly so in the
Trochilidee, but, in so far as that goes as indicating any athnity
between the two groups, it stands for very little.
The external aural aperture in the skull of Arachnothera is
large, and admits of a full view of the interior of the bony ear.
Posteriorly, and to some extent below, this entrance is protected
by a thin scroll of bone, seen in so many other passerine forms,
and very prominent in the Trochilide * # Among some of the
Ceerebide, as, for example, in Diglossa bar itula, these bony parts
of the external ear are remarkably developed ; the aperture upon
either side looks directly to the front, while the bulbous, thin,
and scroll-like wall protecting it above, behind, and below isa
striking feature upon this aspect of the cranium.
Posteriorly, the skull of Avrachnothera ofters but little for
examination beyond what we would find in the skull of any
ordinary passerine bird. The occipital ridge is but faintly defined,
while the supra-occipital prominence is above the average in size,
and especially so in A. longirostris. Thisis also seen in Z’rochilus,
whereas among the Meliphag gidee it is not the case.
Passing to the basis cranii we have tomote the large subcireular
foramen magnum, and the extremely minute occipital condyle,
which is distinctly hemiglobular in form. The basitemporal area
is smooth and convex throughout, presenting at the usual
localities the foramina for the entrance and exit of vessels and
nerves to and from the cranial cavity. The double entrance to
the Eustachian tubes, one to either side, is shielded below by
a very narrow rim of bone. The basi-pre sphenoidal rostrum
presents nothing peculiar, and is thoroughly coéssified anteriorly
with the mesethmoid and the pars plane.
The quadrate possesses a sharp, compressed orbital process, that
in articulation comes in close contact with the cranium. Its
mandibular facet for the lower jaw is double, there being a small
mesial elliptical facet and an outer and larger irregular-shaped
one. A longitudinal groove stands between them. As usual, the
mastoidal articular head is double, and the bone, as a whole, is
highly pneumatic. Apart from the quadrates, all the other
osseous structures at the base of the cranium in Arachnothera are
characterised by extreme slenderness and delicacy of structure.
And, in passing, it may be said that all the articulations,
* Suuretptr, R. W. “Contribution to the Comparative Osteology of the
Trochilide, Caprimulgide, and Cypselide.” P. Z. S. Lond., Dee, 1, 1885, pl. lviii.
fig. De
1909.] OF THE PASSERINE BIRD ARACHNOTHERA MAGNA. 531
especially at the base of the skull, in this genus of birds are
notably small, to the very limits of minuteness. This includes
the articulation of the skull with the spinal column, the quad-
rato-jugal articulation, those with the pterygoids, and to some
extent others.
These last-mentioned bones are very slender, straight, and
rather short, presenting nothing peculiar in their articulations
with the quadrates and palatines. They do not appear to be in
contact with each other in the middle line, though they do
articulate with the sphenoidal rostrum.
The postpalatine portion of either palatine is a delicate scroll
of bone that articulates mesially with its fellow of the opposite
side beneath the presphenoid, while the prepalatine portions are
well apart, straight, and here reduced to a degree of slenderness
rarely met with in birds of this size. A vomer is well-developed,
spatulate in outline, and compressed throughont in the vertical
direction. It is firmly codssified with a palatine upon either side.
Each mawillo-palatine is reduced to the extreme in the matter of
delicacy of structure, being but feebly developed.
Coming to the mandible it is to be observed that it has the
long V-shaped pattern, with a curvature for its anterior two-
thirds corresponding to the curvature of the upper jaw, while its
posterior third is somewhat flexed upon the anterior part of the
bone (see Plate). The rami are very slender and very narrow
from above downward, the structure upon the whole impressing
one with its feebleness. A. magna has the length of the
symphysial portion about equal to the posterior moiety of the
bone; in A. longirostris it is considerably longer, and in this
species, too, the curvature is greater and, if anything, the bone
still weaker. Inferiorly, the “symphysis is smooth, and roundly
convex transversely ; the tomia for this part being sharp. A
small ‘‘ramal vacuity” is present in the mandibles of both these
species, and the free ramal extremities are more or less pneumatic.
There is at each end a small blunt postangular process, otherwise
these ends are practically truncated and concaved behind. The
usual inturned angular processes are present, each having at its
tip or apex the pneumatic foramen found there in so many of the
Class. The coronoid processesare aborted. In A. longirostris the
mandible is 4°8 cm, long and only 2 mm. deep at its deepest
part, about opposite the ramal vacuity.
When normally articulated, the superior and inferior mandibles
in the skull of Arachnothera ave in contact for their entire
lengths. I find nothing to note especially in regard to the
intrinsic ossicles of the internal ear, the siphoniwm, or the
sclerotals of the eye. All are exceedingly passerine in character,
For a representative of this group, however, the hyoid arches
in this species are remarkable. Not only is the glossohyal
greatly elongated to meet the requirements of the feeding-habits
of the bird, but the thyrohyals are similarly produced. The
532 DR. R. W. SHUFELDT ON THE COMPARATIVE OSTEOLOGY [ Apr. 27,
distal ends of the latter run out to hair-like proportions, which
in life curve over the top of the skull, being harboured in the
groove there formed for their reception. This admits of very
considerable extension on the part of the tongue. The cerato-
branchial elements of the thyro-hyals are very long, each being
about half the length, or rather less, than the corresponding
epibranchial. They are more or less straight, and take no part
in the curvature of the posterior ending of this lingual apparatus.
The basihyal is very short, and possesses, distally, a circular tip
for articulation with the glossohyal. A slender, very short,
straight wrohyal is present; the heads of the ceratobranchials
articulating, one on either side, at the junction of the basi- and
urohyal.
A. longirostris has the skeletal parts of its tongue as they exist
in A. magna. The general structure is the same among most of
the Meliphagide; but in that family there exists no marked
elongation of either the glossohyal anteriorly or of the thyro-
hyals behind. In them the lingual apparatus is typically
passerine. Exceptions to this rule, however, exist, and in such
long-billed forms as Acanthorhynchus and some few others the
skeleton of the tongue agrees more or less with what has just
been described for Arachnothera.
Glancing for the moment at the skulls of other species repre-
senting other families at hand, it is to be noted in the skull of
such a bird as Diglossa baritula that in the case of the interorbital
septum it is almost entirely absorbed, a very thin and extremely
narrow piece of bone simply spanning its centre, and the
minutest possible spanlet below is just sufficient to individualize
the two foramina rotunda. The anterior wall of the brain-case
immediately above where these two striplets of bone join is
entirely absent except a very narrow strip just within the orbital
borders. Its occipital condyle is barely any larger than is to be
found in Z'rochilus, and its sphenoidal rostrum is much com-
pressed from side to side. Apteriorly, the rhinal chamber is very
poorly off for bony protection, inasmuch as the elliptical external
narial apertures are very large for the size of the beak: there is
not a vestige of an internasal septum, while the palatal processes
of the premaxillary almost require a lens to see them at all. In
this species the ramal vacuities of the mandible are larger than
we find them in dArachnothera, although the latter is a bird
double its size.
Cinnyris chalybews presents some interesting cranial characters
of its own, for here we find the nasal bones reduced to their very
minimum proportions; the external narial openings are large,
being barely separated above by the culmen. Pars plane are
much reduced in size, and the fronto-interorbital area on the
superior aspect of the skull of this species is notably narrow
transversely. Its mandible is feebly constructed, and the whole
beak considerably decurved.
1909.] OF THE PASSERINE BIRD ARACHNOTHERA MAGNA. 533
In Cereba chloropyga the skull is typically passerine, and in
some respects resembles the skull as found in certain American
Warblers, being quite distinct from what we find in Arachnothera,
to which genus it bears no special aftinity.
So entirely different is the skull in such a species as Prosthema-
dera novee-hollandie of New Zealand, a bird placed among the
Meliphagide, that a separate description would be required to
give an account of it. Here the masals are very broad antero-
poster lorly, and each is pierced by a central foramen, an unusual
character. Then the pars plane are very thick from before
backwards, and a longitudinal groove marks the external aspect
of each.
In not a few particulars Acanthogenys rufigularis of Australia
is a Meliphagidine species with a skull not at all unlike what we
find in the species of Acanthorhynchus, and these forms are more
or less nearly related. Acanthogenys has the broad nasals, each
pierced by the small central foramen, and there are several
other points in the two skulls of more or less close agreement.
But such representatives of the Meliphagidee have no special
relationship with the Coerebide, and even less with the typical
Nectarinude. Judging from the skulls alone, it is not difficult to
recognize the more or less close relationship existing among the
species I have before me of the genera Hntomyza, Acanihogenys,
and Prosthemadera, all of which present characters in this part
of the skeleton quite different from anything we find in Arachno-
thera, and surely offer no skull-characters at all approaching any
of the Trochilidee.
I have made no attempt to either study or compare the ossifica-
tions presented on the part of the trachez in any of these birds.
From superficial examination only, I would say that although
generic and family differences-are easily to be seen in these parts,
yet at the same time no very striking departures are to be noticed
from the general passerine character in any of them.
Having then compared the morphological characters of the
skull and the associated osseous structures in such species as
there are at hand representing the families Coerebide, Necta-
riniide, Certhiide, and Meliphagide, and these characters with
the corresponding ones in the skull of Z'rochilus, it is clear that,
in so far as this part of the skeleton is concerned, these four
passerine families are a very long way removed from the Super-
suborder Zrochiliformes, and this is no more than one would
naturally expect to find.
My views upon the position in the system of the four above-
named families have already been published, and I see no special
reason for changing them*. From this point on any further
comparison of the skeletons of these birds with the osteology of
* Suureipt, R. W. “‘ An Arrangement of the Families and the Higher Groups
of Birds.” Amer. Nat. vol. xxxviii. nos. 455-456, Boston, Nov.—Dec. 1904, pp. 835-
856.
534 DR. R.W.SHUFELDI ON THE COMPARATIVE OSTEOLOGY [ Apr. 27,
the Humming-birds would be quite unavailing. In all respects
it is very different. I shall proceed, then, to complete this
account with a description of the remainder of the axial skeleton
and the skeleton of the limbs in Arachnothera, comparing the
principal characters with those presented by the skeletons of the
other families above-named.
REMAINDER OF THE AXIAL SKELETON.—So far as the vertebre
are concerned between the skull and the pelvic sacrum, the
several families of birds here being considered are all strictly
passerine in character. I have counted and compared them in a
representative of each family, and am satisfied that these bones
offer nothing worthy of an extended and detailed description.
To be sure, we find some differences in form among the various
species, genera, and families, but such characters are of but slight
importance, and in all instances the variations are no more
marked than those which obtain among more or less nearly related
passerine birds in the avifauna of any country of great extent,
as, for example, in Stalia, Wimus, and American Warblers, as
compared with the Passeres of the Pacific Coast region*. Both
Arachnothera magna and A. longirostris possess 19 vertebre
between the skull and the pelvic sacrum; the first twelve are
true cervical vertebree without free ribs. The 13th and 14th are
also cervicals, the first supporting a pair of very small free ribs,
and the last a far better developed pair, which are likewise free
and without wnciform processes. The next five are true dorsals
and possess these appendages, and also connect with the sternum
through the intervention of costal ribs. We also find a very
delicate pair of ribs attached to the leading vertebra of the
sacrum, the sternal ribs of which are ‘floating ribs.” Dorsal
vertebre have well-developed neural spines which are in contact
with each other anteriorly and posteriorly, the ultimate dorsals
being lacking in hemal spines.
There are seven free caudal vertebre in the skeleton of the tail
in A. magna, in addition toa rather large pygostyle. The form
assumed by the latter is well-shown in the Plate illustrating this
paper. In the family Meliphagide there appear to be but six
caudal vertebre and the pygostyle, and this is likewise the case
among the Ccerebide, as, for example, in Cyanerpes cyanea.
However, we know that even in the same species these caudal
vertebra may vary to the extent of one, or may be two; they are
not constant even 1n our own species.
As well as can be made out from an adult skeleton, or rather
a skeleton from an adult individual, it would appear that there
are eleven vertebre fused together to form the pelvic sacrum in
* SuuretpT, R. W. “Contributions to the Comparative Osteology of the
Families of North American Passeres.” Jour. Morph. vol. ui. no. 1, Boston,
June 1889, pp. 81-114, pls. v. & vi. A large number of the passerine birds of the
United States are osteologically compared in this paner.
1909.] OF THE PASSERINE BIRD ARACHNOTHERA MAGNA, 535
A. magna, and they present the usual passerine characters. Only
tei, it would seem, are thus fused together among the Cerebide,
but in these birds the pelvis is relatively shorter and wider than
it is, asa rule, among the Nectariniidee. The Meliphagide have
eleven—that is, in the species at hand, though Acanthogenys
rufiguiaris may be an exception and possess only ten; in any
case, an extra vertebra in the sacrum may account for one less in
the caudal series.
Among all small, ordinary, and more or less typical Passeres all
over the world where they may occur, we meet with but little
variety in the form of the pelvis. Its passerine characters are
very uniform. Some birds of the group have it rather narrow
and deep ; in others it is wider and more compressed from above
downward, with the pubic elements far apart—but the general
characters remain the same. In 4A. magna it is of the narrow
and deep variety, with the iio-neuwral canals open for their entire
lengths, and the “sacral crista” standing between them very
prominent. On the postacetabular area parial foramina occur
among the diapophyses of the ultimate sacral vertebrae. Anteriorly
the ilia are truncated from their mesial angles, backward.
Posterior to an acetabulum, on a side view, we note the large
ischiadic foramen; a small, circular obturator foramen, which is
barely separated from the large tendinal vacuity. Behind this
last the ischium dips down, as usual in Passeres, to meet the
pubis, or pubic style, near its distal termination. The post-
acetabular, external free margin of the ilium to some extent
overhangs the ischiadie foramen, while the preacetabular part
of one of these bones is hollowed externally throughout its
extent. The pelvis exhibits specific differences in 4. longirostris,
where the postsacral foramina are large, and the internal iliac
margins in the postacetabular region do not codssify with the
sacrum.
Among the Meliphagide, we find that in Entomyza cyanotis
the ilio-neural canals or “grooves” remain open only anteriorly,
and this is the case with other species of this family. On
lateral view of this pelvis all the openings are very large, the
osseous side of the bone here being reduced to the minimum
thereby. This statement refers also to the large circular coty-
loid ring (acetabulum) and the mergence of the obturator
foramen with the tendinal vacuity. In Cyanerpes cyanea the
essential characters are the same, but the pelvis, as a whole,
is broader and flatter than it is in Arachnothera, coming in
this respect nearer some of the Nectariniide, as, for example,
Anthreptes malaccensis.
Passing to the shoulder-girdle and sternum, we find all this part
of the skeleton very distinctly passerine in character, the matter
of size of the bones often being all there is to differentiate them
with respect to the species they belong to: as, for example, the os
furcula of such a species as Prosthemadera novee-hollandice is, as
536 DR. R. W.SHUFELDT ON THE COMPARATIVE OSTEOLOGY [ Apr. 27,
we would expect, a larger bone than the os furcaula of Arachno-
thera magna, yet the characters are identically the same, so much
so that were the bone in the latter brought up to the size of the
one in the former, I very much doubt that any ornithotomist
could, with certainty, tell them apart. Indeed, the os furcula
possesses in all the passerine birds here under examination
the same form and characters. When we say it is U-shaped
in outline, has a large, transversely compressed, and up-
turned hypocleidium, slender limbs, and expanded clavicular
heads, we have said about all there is to be said in regard
to it.
All these birds have coracoids and scapule of the shoulder-girdle
very much alike indeed—that is, apart from the matter of size.
The morphological variations are very insignificant and offer
little or nothing of taxonomic value. It is interesting to note,
however, that irrespective of the length or form of the beak, we
find in all the Meliphagide that the lower external angle of the
expanded part of a coracoid is produced outwards as a distinct,
flattened process, best seen in the short-billed species, though
also well-marked in Acanthorhynchus, which Dr. Richmond informs
me is a genus belonging to the family Meliphagide. Now, in
A rachno® -ra@, and in all the Nectar iniide and Ccerebide at hand,
that anjle of the coracoid is more or less truncated, but whether
this points correctly to any Sassi relationships of the families
named, it would be difficult to say *
Representatives of all these niles at hand, as in all true
Passeres I believe, have at each shoulder-jomt an os hwmero-
scapulare, and it varies but very little in size and form in the
species examined. When we come to examine the sternum of
Arachnothera magna and compare it with the sterna of other
species of Nectariniide, and with the other bird-fornis enumerated
in this paper, we once more realize that this part of the skeleton
is likewise all passerine in its morphology, presenting only a few
slight differences for the various species. Still, apart from the
variations in size, these differences are more or less constant, and
in any case anrisialihle with respect to the sternum Be the
species possessing them—that is, the species which have thus far
been named in this paper. The sternum in all exhibits a number
of characters which all the sterna present in common, as the
large, upturned trihedral manubrium, with its bifurcated free
extremity; the lofty costal processes with the hemapophysial
facets on the posterior borders; the marked concavity of the
* Without going into details. and judging from the skeletons alone, I would
remark that the Australian genus of birds named Acanthor hynchus which have been
referred to the Meliphagide possess skeletal characters which in the main agree
better with the corresponding ones in Arachnothera than they do with any ot “the
same characters as seen in the short-billed Meliphagids. But_two species of
Acanthorhynchus are known to me, A. fenuirostris and A. superciliosus, and these
I have never had the opportunity of comparing in the flesh with the meliphagidine
birds of Australia and New Zealand.
1909.| ‘OF THE PASSERINE BIRD ARACHNOTHERA MAGNA. 53
dorsal aspect; the pair of “notches” in the xiphoidal extremity
giving rise to an outer xiphoidal process upon either side; the
deep “keel” with its prominent carinal angle, anteriorly; and,
finally, the fact that the bone is more or less pneumatic. The
coracoids never decussate in their sternal beds, though in some
species they almost appear to do so. Again, the hypocleidium of
the furcula never comes in contact with the manubrial process of
the sternum, though in some species the approach is extremely
close (Hntomyza cyanotis). So much for the general characters,
and Arachnothera has them all to perfection ; and there are a few
minute pneumatic foramina in the middle line on the dorsal
aspect of the bone, anteriorly, in this species. If we designate the
length of the sternum by a line extending from the apex of the
earinal angle of the keel to the mid-xiphoidal point posteriorly
where the keel terminates, then this distance in A. magna
measures 2°3 centimetres. This same measurement will be
applied in the case of the sterna of the other species—as, for
example, in Arachnothera longirostris, the length of the sternum
is but 1°5 ems., and we find the body of the bone flatter, the
xiphoidal notches comparatively deeper, the carinal a gle not so
acute, and the keel of the manubrium conspicuous and vroduced
well down upon the anterior border of the sternal cari. . The
pneumatic foramina are scarce and in the same localit,. In
Leptocoma grayi the sternum has a length of but 1:3 cms.
In Cyanerpes cyanea, as representing the Coerebide, it measures
1:7 ems., and here the bone is very thin and delicately constructed,
with a small manubrium; deep notches and dilated extremities
to the xiphoidal processes. These characters do not apply to the
sternum of Cwreba chloropyga, another of the Cerebide, a species
having the bone only 1:2 ems. long.
Anthreptes malaccensis has a sternum which is the counterpart
of that bone in A. magna, only it is much smaller, having a length
of but 1:4 cms.
Among the larger forms of the Meliphagidee we meet with
characters in the sternum that are absolutely diagnostic, as, for
example, there is no such a thing as mistaking the species had we
but this bone to help us in such a bird as Acanthogenys rufigularis.
Here, although it is of the usual passerine type, it is peculiar in
having the anterior border of the sternal body very much
thickened and rounded; this thickening is continued across the
base of either costal process and up on to the mesio-posterior
margin of the same. Such a thickening also defines the limits
laterally of a deep mesio-longitudinal groove, deepest anteriorly,
that 1s found upon the dorsal aspect of the body of the bone.
For its anterior.moiety, thickly crowded together at the bottom
of this groove, we find some thirty or forty pneumatic foraminal
openings. Between the coracoidal grooves there is another
single pneumatic foramen, and the external angles of the mid-
xiphoidal prolongation are pronounced. The carinal angle is
Proc, Zoou. Soc.—1909, No. XXXV. 35
538 DR. R. W. SHUFELD? ON THE COMPARATIVE OSTEOLOGY | Apr. 27,
not prominent and is to some extent slightly rounded off. Thus
it will be observed that this meliphagidine species possesses a
very characteristic sternum, quite different from the bone in
Arachnothera.
In Entomyza cyanotis the sternum, of course, is somewhat
larger, and in it the anterior border of the body is thickened
and the mesial groove with its pneumatic foramina on the
dorsal aspect is confined to the anterior third of the bone.
The carinal angle is sharp again and rather prominent, while the
most distinctive character of this sternum is the unusually short
manubrial process, though its bifurcations are conspicuous. In
this bird I should have noticed above that the coracoids are very
long and particularly slender.
In Prosthemadera nove-hollandie the sternum is likewise
distinctive, in that the above-described mesial groove on the dorsal
aspect is absent, and the pneumatic foramina in that locality are
reduced to a very few minute and hardly noticeable ones. Very
little thickening of the anterior sternal border is to be observed,
and the carinal angle is not produced anteriorly. As compared
with other Meliphagide, however, the sternum of this species
may be easily recognized by the remarkably thickened free
anterior border of the manubrium, which is extended more than
halfway down upon the anterior border of the keel. Then,
posteriorly, the external lateral xiphoidal processes are unusually
slender and long. Distally, on either side, they almost touch the
greatly produced outer angle of the mid-xiphoidal prolongation,
thus giving the much elongated “notch” upon either side the
appearance of a foramen.
Posteriorly the keel terminates in a conspicuous triangular
area, and, as usual, there are jive hemapophysial facets on the
posterior border of either costal process.
Entomyza cyanotis has the sternum 3°2; <Acanthogenys rufi-
gularis 2°6; and Prosthemadera nove-hollandie 3:1 centimetres
long.
So far as any characters go, the sternum of the last-named
species is most like the bone in dArachnothera magna, and this
seems to be true with respect to some other parts of its skeleton.
Whether this circumstance carries with it the fact that these
two species are more nearly affined than either of the other two
meliphagidines just named remains to be seen. It would require
more extensive and varied material, I must believe, to be certain
of such a point as this.
OF THE APPENDICULAR SKELETON: THE PEcroRAL Lims.—
With great care I have examined and compared the bones of
both the pectoral and pelvic limbs in the birds named in the
list presented in the first part of this paper. This part of the
osseous system has likewise been studied by me in a very large
number of passerine species from various parts of the world. The
1909.] OF THE PASSERINE BIRD ARACHNOTHERA MAGNA, 539
forms here being examined are from Mexico, South America,
South Africa, Asia, East India, New Zealand, and Australia,
or, in other words, very widely separated countries, yet there is
no mistaking any of the characters of the skeleton of the limbs in
any of the birds,—they are all distinctly passerine, we might say
“typically” passerine, had there ever been selected as a reference
standard a species anywhere, the skeleton of which was to be
universally considered as the type in this respect for all Passeres.
It would be interesting to know what species would be selected
upon a consensus of opinion of ornithotomists everywhere, on
this point—WMerula perhaps. Typically passerine o1 otherwise,
however, one would be surprised to note the variations in form in
some of these bones among these oscine honey-birds, creepers, and
their kin. ‘To appreciate this to the full extent, the skeleton of
the limbs of the several species would have to be brought up,
either in drawings or plastic models, to many times the size
of the original, and then compared one with another. ‘Take
the humerus, for example: had we models of this bone from
all the species in our list each twenty centimetres long, and
the other proportions correctly reproduced, we should be sur-
prised at the differences which exist in them, which by this
means could easily be recognized at a glance, but which other-
wise must be studied by passing from one to the other with a
good lens to assist you.
In Arachnothera magna the humerus has a length of 2°2 ems.,
and its smooth shaft is nearly straight, presenting hardly any of
the usual sigmoidal curve from whatever point we may view the
bone. In form it is subcylindrical, being somewhat compressed,
from its anconal aspect palmad. The radial crest is short and net
very prominent, while the ulnar tuberosity is conspicuous, and
the pnewmatic fossa surrounded by a distinct raised margin, which
between this concavity and the head of the bone on the anconal
side is raised into a thin and projecting process. What is most
peculiar is that the head of the humerus is hollow and the
entrance to the concavity is seen on the anconal side, next to the
pneumatic fossa, being separated from it by the aforesaid thin
projecting process. A few small, scattered, pneumatic foramina
are found at the bottom of the fossa, where they usually occur.
At the distal end of the bone we find the oblique and ulnar
tubercles much produced with the olecranon fossa circumscribed
and well marked. It is confined to the ulnar side of the shaft.
The ecte- and entepicondylar processes are both distinct though
not large apophyses. In all these birds there is to be found a /ree
sesamoid at the elbow-joint.
Arachnothera longirostris has a humerus resembling the bone in
A. magna, but in it the caput humeri is not hollowed out in the
manner described above. In both species there is a distinct and
circumscribed pit at the distal end of the shaft on the palmar
surface right next to the ulnar tuberosity. This humerus has a
540 DR. R. W. SHUFELDY ON THE COMPARATIVE OSTEOLOGY [ Apr. 27,
length of 1-7 ems. Considering its size, the proximal end of the
bone is expanded in Leptocoma gray, and the processes at the
distal extremity conspicuous; otherwise the characters are quite
identical with those in the humerus of 4. magna. It has alength
of 1-3 ems.
Cinnyris chalybeus has the caput humeri but very slightly
hollowed out, and the bone has almost the appearance of being
non-pneumatic. In any event, by the aid of a strong lens, I
failed to find pneumatic foramina at the base of the fossa.
The radial crest is considerably aborted, and a long shallow
notch cuts out a portion of its free continuity. The bone is
1:2 ems. long.
Among the Cerebide, Diglossa baritula has a non-pneumatic
humerus, measuring in length 1:2 cms. Its distal processes and
tuberosities are made conspicuous by their distinctness—clean-
cut and produced, as it were. There is not a little hollowing out
of the head of the bone next to the pneumatic fossa, the concavity
being fairly well separated from the latter by a thin osseous
partition, as in 4. magna. These two cavities practically merge
in such a form as Anthreptes malaccensis, and the partition
between them in other species, though well produced, has with-
drawn toward the proximal end of the bone. It has a length of
1-6 cms. and appears to be non-pneumatie.
In Cyanerpes cyanea this mergence of the pneumatic fossa and
the very profound excavation of the caput humeri is practically
complete—the merest vestige of an osseous partition existing
between them. The radial crest is very short ; and this humerus,
too, ERE S to be non-pneumatic or very largely so. It hasa
length of 1:5 ems. Without particularly examining this bone in
other Nectariniids, Ceerebide, or Certhiide though it may be
said with respect to the latter that in Climacteris scandens the
humerus presents very different characters, for in it the caput
humerit is not hollowed out at all, the pneumatic fossa is very
open, the foramina large, the entire bone pneumatic, and the
shaft somewhat curved (length 2 ems. )—we may pass toa study
of the bone in the Meliphagide. Here in Acanthorhg ynchus,
Entomyza cyanotis, and Acanthogenys rutigularis the caput humeri
never exhibits the slightest indication of any hollowing out
or any cavity leading in to it next to the pneumatic fossa.
The latter is large, deep, with its free margin much thickened,
and the pneumatic foramina at its bottom big and more or less
numerous, as the bone in these birds is ‘highly pneumatic.
The subeylindrical shaft displays to same degree the sigmoidal
curve and in some species is stout for its length, as in
Prosthemadera nove-hollandie. he radial crest is short and
not very prominent, while the tubercles and processes at the
distal end are so.
Passing to the bones of the antibrachium, carpus, and manus
we find that they present among these families, including
1909. | OF THE PASSERINE BIRD ARACHNODIMERA MAGNA, HAI
Arachnothera magna, fewer distinctive characters than do the
humeri for the species examined. All are essentially passerine
and more or less typically so. Radius and ulna of the fore-
arm are invariably straight, and present the characters common
to the group. On the shaft of the ulna the papille for the quill-
butts of the secondary feathers of the wing may be very well
pronounced (Hntomyza) or entirely absent (Arachnothera, Pro-
sthemadera, and most others).
The middle metacarpal is always produced beyond the main
bone of the metacarpus—the index metacarpal,—and I have never
met with any claws on any of the terminal phalanges. There is
always present a triangular, flattened process at the proximal end,
ulnar side, outer aspect of the index metacarpal which is directed
backward and rests flat against the proximal end of mid-
metacarpal.
Radiale and ulnari of the wrist are invariably well-developed
and present the usual passerine characters.
THe Petvic Lims.—Upon comparing the bones entering into
this extremity in Arachnothera magna with the corresponding
ones as we find them in the legs of the cerebidine, certhidine, and
meliphagidine species at hand, it is to be observed that the
characters upon the whole are more uniform than they are for
the bones of the pectoral limb. Osteologically, the limb is of a
strictly passerine type throughout, and when compared, bone for
bone, with the limb in any of the non-passerine suborders, presents
more or fewer differences. As a rule, in the former the long
bones of the thigh, leg, and metatarsus are quite straight, and in
the case of the femur and tibio-tarsus have cylindrical shafts.
Entomyza cyanotis offers an exception in the case of the latter
bone, for instead of its shaft being entirely straight it is seen to
curve away from the fibula above the fibular ridge and approach
it again proximally to articulate with the latter and the
femoral condyles. This condition is not usually seen or is much
less marked among other meliphagidine species. As in the case
of the bones of the antibrachium and manus, all the bones of
the pelvic limb are non-pneumatic, and the nutrient foramina
that enter them are very minute.
In the femur the semi-globular head is sessile, and the pit for
the ligamentum teres generally very feebly marked. Distally, the
summit of the bone includes a part of the head, the great trochanter,
and the valley between the two, thus forming one surface, which
is articular, and lies in a plane to which the longitudinal axis of
the shaft of the bone is perpendicular. Distally, the condylar end
of the femur is well-developed and presents the usual passerine
characters. The depth of the rotular channel varies in different
species and families, being rather deep in Arachnothera and
generally shallower in the Meliphagidee.
As in the majority, if not in all, oscine passeres, the species
542. -DR. R. W. SHUFELDY’ ON THE COMPARATIVE OSTEOLOGY [ Apr. 27,
here under examination possess a well-developed osseous patella.
As we would naturally expect, it varies in size for the species, but
very little with respect to form. In Arachnothera magna it is of
a cordate outline, with the much rounded apex below, convex
anteriorly, and decidedly concave on its upper and _ posterior
aspects. Posteriorly, the concavity is double, the surface between
being intended to accommodate itself to the femoral condyles. In
Prosthemadera nove-hollandie the patella has a transverse
diameter of half a centimetre, and is two millimetres deep at the
centre.
Tibio-tarsus supports the usual pro- and ectocnemial processes,
and these have their common passerine form, varying but slightly
for any of the species here being examined. In all cases they rise
but slightly above the summit of the bone; they extend only a
very short distance down the shaft, and both are always turned
slightly fibulawards. Their antero-inferior angles may be sharp-
pointed (Arachnothera, Cereba, and Diglossa) or they may be rounded
off as in the Meliphagide. They are very conspicuous and far
apart, and produced almost directly to the front in Climacteris
scandens. Always feebly developed, the fibula is but rarely pro-
duced beyond, or much beyond, the fibular ridge on the tibio-tarsus
in articulation. It is a weak bone, of but slight importance in all
small passerine forms.
Tarso-metatarsus presents nothing of marked departure from
the ordinary form of the bone among all small passerine birds.
Its hypotarsus is always reduced to a small subeubical apophysis
vertically pierced for the passage of tendons. The shaft is
always more or less flat anteriorly, and longitudinally grooved
behind. The three distal trochlear processes are found in nearly
the same transverse plane, with a large, free accessory meta-
tarsal directed backward.
As to the skeleton of the pes it is purely passerine in its mor-
phology, with the joints of the toes on the usual plan. All of
these species have the ungual joints large and much curved,
especially is this the case with respect to the ungual phalanx
of the hallua: in the larger species of the Meliphagide.
Ossification in these birds, and probably in others, may
normally extend to some of the tendons of the muscles at the
back of the tarso-metatarsus between the hypotarsus and the
accessory metatarsal of the hallux, which is markedly the case in
Entomyza.
CoNCLUSIONS.
It is clear from what has been brought out in this paper that
none of the species of birds here osteologically passed in review,
employing Arachnothera magna as the type, has any especial
affinity with the Trochili, Even the morphology of the tongue in
quite a number of the forms has no significance when taken in
1909.] OF THE PASSERINE BIRD ARACHNOTUERA MAGNA. 543
connection with the rest of the anatomical structure, which in
the Humming-birds is so decidedly non-passerine, while in all
the families here investigated the osteology at least points most
distinctly to the Passeriformes.
There are a few faint cranial resemblances to be seen upon
comparing that part of the skeleton of a Humming-bird with
the corresponding structures in certain Corebide, or even
Nectariniide, but they amount only to resemblances and no
more, ‘hese are interesting, but they have no bearing whatever
upon affinities.
As to the probable relations of these several families—the
Nectariniide, Corebide, Certhiide, and Meliphagidee—to each
other and to other groups, we are confronted with an entirely
different question, and one by no means easy of settlement.
In my paper on “An Arrangement of the Families and the
Higher Groups of Birds” my provisional opinion has already been
presented. Therethe Meliphagide stand between the Nectariniide
and Certhiidee—the three families being kept together and placed
between the Diceeidee and Mniotiltide. From these the Corebide
are well separated, being placed between the Drepanide and
Tanagride, with several families intervening, including the Larks
and Finches. I do not know that this opinion has in any way
been shaken or disturbed by the examination of the material
Mr. Beddard has so kindly placed at my disposal ; still before my
convictions come to be any firmer or more settled, I would prefer
to investigate the anatomy of a great many more species of
families evidently related, than I have up to the present time.
Were I to propose a change in my arrangement I would say
perhaps that the Cerebide should hold a place in closer relation-
ship to the Creepers and Warblers, which I am rather inclined to
think that they do. As a family, however, the Carebide are
undoubtedly very distinct, certainly from the Nectariniid, here
made to include the former family Cinnyride. The examination
of the skeleton of such a form as Diglossa baritula would be quite
sufficient to establish that fact, and this has been done in the
present contribution. A few more Sun-birds and Honey-suckers
should be examined, especially of the genera Drepanis and
Meliphaga, which I have not as yet seen. Cinnyris is a genus of
the Nectariniide, though not typical of that family. I have in
former years examined many of the Sittidee—the Honey-creepers
(Cereba) are quite distinct from them.
Climacteris scandens as a representative of the Certhiid
clearly points to the fact of the distinctness of that family,
but I should like to examine the skeletons of other species
of Wall-creepers.
A far greater number of the Meliphagide need investigation,
and the genera here examined do not point, in so far as their
skeletons go, to any very close relationship with the Nectariniide,
as exemplified in the genus Arachnothera and others. Especially
p44 OSTEOLOGY OF ARACHNOTHERA MAGNA. Ayer an.
does this required research apply to the subfamiles J/yzomeline,
Melithreptine, and Meliphagine. Possibly the representatives of
some of these are more nearly affined to such a species as
Arachnothera magna than some we have seen. In any event my
labour will not have been in vain if what has here been brought
out with respect to the osteology of the four meliphagidine genera
examined proves to be of any assistance in that direction.
EXPLANATION OF PLATE LXVIII.
Left lateral view of the skeleton of Arachnothera magna. Adult. Natural size.
Right ulna fractured at its proximal third. Preparation partly ligamentous ;
podothece not entirely removed. Reproduced from photograph of the specimen
by the author.
Abantis
efulensis, 407, 409.
elegantula, 407, 408,
409.
qa, 407, 408, 413.
lucretia, 407, 409, 413.
Acallopistes, 407.
Acanthogenys, 528.
rufigularis, 627, 629,
533, 535, 537, 538,
540,
Acantholeberis
curvirostris, 329.
Acanthorhynchus
sp., 927, 528, 529, 532.
superciliosus, 536.
tenuirostris, 536.
Acleros
kasai, 410, 413.
leucopyga, 410.
sparsum, 407, 410, 413.
Acmea, 257.
testudinalis, 242, 248.
virgined, 242.
Acomys
scloust, 121.
Acroperus
harpe, 330.
Aerydium
sp., 157.
japonicum, 157.
Acta
fossulata, 159.
.Epyceros
melampus, 125.
Alschna, 284.
/Ethus
pygmaeus, 153.
Agapornis
nigrigenis, 130.
Agrion
puella, 256, 266, 276,
217, 285.
INDEX.
Agrion
pulchellum, 255, 2857,
258, 262, 263, 266,
267, 269, 272, 278,
216, 277, 2895.
Aleochara
sp., lol.
Aleurobius
Jarine, 327.
Allacta
noctulata, 156.
obtusata, 156.
Alona
affinis, 330.
quadrangularis, 330.
Alonella
excisa, 329, 330.
nana, 329.
Alonopsis
elongata, 330.
Alpheus
strenwus, 160.
Ameiva
surinamensis, 227.
Amoeba, 309.
Amphiascus, 40.
Amphioxus, 203.
Anax
guttatus, 155, 156.
Andronymus
leander, 407.
Anisolabis
annulipes, 156.
Anoa
depressicornis, 90.
Anomala
sp., Lol.
Anous
stolidus, 137.
Anser
cinerea, 840.
Anthrax
sp., 150.
Proc. Zoot. Soc.—1909, No. XXXVI.
Anthreptes, 528.
malaccensis, 527, 53d,
537, 540.
Anthus
bertheloti, 370.
bosniaskii, 369, 370.
Antilocapra, 174, 175,
V7, 179, V8, 182;
185, 186, 190, 197.
americana, 160, 172,
173, 178, 183, 197.
Antilope
dorcas, 340.
Aonyx
capensis, 119.
Apatura, 282.
Apaustus
abure, 411.
Aprosmictus
chloropterus, 130.
Apteryx, 224.
Arachnothera
longirostris, 527, 528
ays) Sh), GRil., Bay,
584, 839, 937,
539.
magna, D27-544,
Aradus
membranaceus, 15+.
Archispirostreptus
eristulatus, 414.
pyrocephalus, 414.
Arctictis, 89, 479.
binturong, 90, 386.
Argas
persicus, 327.
Argulus, 11, 12.
Arion
ater, 328.
empiricorum, 623 329
332.
Artholexis
hollandi, 407.
36
XXV1
Artholexis
melichroptera, 407, 408,
413.
Artopéetes, gen.
422, 473, 476.
pryeri, 473, 474, 476.
Artopoia, 473.
Arvicanthis
dorsalis, 121.
pumilio dilectus, 121.
pumelus, 520.
Ascaris
compar, 364.
Suniculus, 365.
gallopavonis, 365.
gibbosa, 365.
znjlexa, 365.
lagopodis, 364.
mucronota, 340.
papillosa, 364.
perspicillum, 365.
vesicularis, 364.
Aspergillus
fumigatus, 128.
niger, 128.
Ateles, 86.
Attheyella
cuspidata, 330.
eschokkei, 330.
Aulacodus
swindernianus, 516.
Aurelia
aurita, 78-81.
cerulea, 80.
— colpoda, 80.
— cruciata, 80.
— japonica, 80.
noy.,
Babyrussa, 1935, 196,
197.
alfurus, 160, 192, 194,
195.
Balena
australis, 95.
glacialis, 91-98.
Balzenoptera
borealis, 92, 96.
physalus, 96.
Balistes
sp., L44.
rectangulus, 406.
Bandicota
setifera, 391.
Baoris
arela, 412.
statirides, 412.
Bassaricyon
alleni, 492.
Bassaris
astutus, 90.
INDEX.
Bavia
sp., 158.
Belemnitella
mucronata, 304, 306.
Belideus
sciureus, 497.
Bernicla
brenta, 227.
Bibio
sp., 361.
Birgus
latro, 136, 159.
Bison
americanus, 308.
Blatella
germanica, 156.
Blennius
atrocinctus, 405, 406.
nativitatis, 404, 406.
Bodianus
Fulvus, 131.
punctatus, 131.
ruber, 131.
Boeckella, 33.
Bombyx
mort, 284.
Bonasia
sylvestris,
368.
Boreophausia
tnermis, 97.
Bosmina
longirostris, 330.
Bothria, 473.
Bothrinia, nom.
425, 473.
chennelliz, 446, 473.
neoulosa, 472, 476.
Bothriocephalus
sp., 367, 368.
Botria, 473.
Brachyrhynechus
membranaceus, 154.
Bradypus
tridactylus, 523.
Brotula
multibarbata, 406.
Bubalus
caffer, 124.
364, 367,
nov.,
Ceenides
cerymica, 407.
dacena, 407.
tlerda, 407.
kanguensis, 407.
lacida, 407.
laterculus, 407.
leonora, 407.
zaremba, 407.
Calappa
hepatica, 159.
Caleinus
herbstiz, 159.
Callithrix, 86.
Callopistes
maculatus, 227.
Caloeyelus
annularis, 153.
Calodium
caudinflatum, 345.
Camelus
dromedarius, 340.
Candona
candida, 380.
Caninia
cornubovis, 285.
cornucopie, 285, 286,
>
Canis
adustus, 119.
mesomelas, 118.
Capra
falconer?, 130.
Capromys, 89.
pilorides, 90.
Caracanthus
unipinna, 404.
Cardisoma
hirtipes, 159.
Cariama, 227.
Carpilius
maculatus, 159.
Carpophaga
whartoni, 102, 137.
Carupa
leviuscula, 159.
Cassiopea, 81.
Castalius, 421.
catreus, 476.
ethion, 423, 475, 476.
Cateptria
sp., 147.
Cebus, 86.
Celenorhinus
nigropunctata, 407,
13.
rutilans, 407.
Celastrina, 419.
Centetes, 510, 512, 513,
525, 526.
ecaudatus, 508, 411.
Cephalophus, 160, 197.
abyssinicus, 189, 190.
dorsalis, 181, 182, 188,
197.
grimmi, 123.
maxwelli, 181, 182,
183, 187, 197.
melanorrhous, 181,
182.
Oephonodes
hylas, 147.
Ceratophyllus
gallinule, 325, 384.
garet, 325,
(Lrichopsylla)
steadi, 325.
Ceratrichia
dimidiata, 407.
ialemia, 413.
nothus, 413.
paucipunctata, 412.
Cercaria
trigonocerca, 328.
Cercariacum
limacis, 328.
Cercocebus *
auratus, 373.
JSuliginosus, 87, 372.
Cercoleptes, 89.
caudivolvulus, 90, 488,
490.
Cercopithecus, 86.
Suliginosus, 87.
pygerythrus, 83, 114,
ruber, 86.
sabeus, 85.
Ceresium
nigrum, 159.
quadrimaculatum, 153,
simplex, 152.
Ceropria
induta, 152.
Cervicapra
arundinum, 123.
Cervulus, 190.
muntjac, 90.
Cervus, 174, 191.
dama, 190.
hippelaphus, 136.
muntjac, 136.
murrayt, 136.
Cheerocampa
vigil, 148.
Chameleo, 227.
Chelmo
longirostris, 403.
Chelone
imbricata, 148.
mydas, 143.
Chilodactylus, 404.
Chinchilla
lanigera, 516.
Chionis
alba, 1.
Chiromys, 522.
madagascariensis, 514,
Chironemus, 404.
Chiropodomys
anna, 890.
gliroides, 390,
new-
INDEX.
Chiropodomys
major, 390.
niadis, 390.
peguensis, 390.
penicillatus, 390.
pusillus, 390.
Chloéon, 280.
dimidiatum, 284.
Chloridea
assulta, 146.
obsoleta, 146.
Chloroclystis
tenuilinea, 148.
Choerephon
plicatus, 383.
Choleepus
didactylus, 523.
hoffmanit, 90.
Choristoneura, 408.
Chrysolophus
(Phasianus) pictus,
346
Chrysothrix, 86.
sciureus, 522.
Chydorus
sphericus, 829, 380.
Cicinnurus
regius, 414.
Cimex
viridula, 153.
Cinclus
aquaticus, 325,
Cinnyris, 528, 543.
chalybeus, 527,
532, 540.
Cireus
ranivorus, 414.
Cirphis
loreyi, 146.
Cirrhites
murrayi, 404, 406.
Cletodes, 46.
Clibanarius
corallinus, 159.
Climacteris, 528.
scandens, 527, 529, 540,
542,
Cleeotis
percivali, 113, 115.
Clupea
alosa, 201.
Coccidium
avium, 309.
Coccinella
transversalis, 151.
Coccothraustes
vulgaris, 325.
Ccelogenys, 185.
Ceelops
bernsteini, 379.
frithii, 879.
529,
XXVil
Ccelops
robinsont, 379.
Coenobita
clypeatus, 159,
perlatus, 159.
rugosus, 159.
Ceereba, 528, 542.
chloropyga, 527, 583
587.
Coleophora
cespetitiella, 332,
glaucicolella, 332.
Columba
palumbus, 325.
Connochzetes
taurinus, 122,
Conocephaloides
sobrinus, 157.
Corvus, 370.
Corynetes
rufipes, 151.
Coturnix
communis, 346,
Crocidolomia
binotalis, 148.
Crocidura
sp., 116, 384.
JSuliginosa, 102,
Orocodilus
palustris, 280.
Crossarchus, 489,
fasciatus, 480, 487,
Orypsirhina
varians, 130.
Cryptoccenia, 292,
Oryptoprecta
Jerox, 499.
Ctenoplana
korotneffii, 80.
Cuon
javanicus, 386,
Cupido, 473,
Cuscus
sp., 497.
Cyanerpes, 528.
cyaned, 527, 584, 535,
537, 540.
Oyaniris, 419, 471.
Cyathophora
pratti, 292.
Cycloeypris
globosa, 330,
serena, dol.
Cyclopides
leucopyga, 41U.
Cyclops, 49, 67.
agilis, 59.
aguloides, 59, 68, 69,
76.
albidus, 53, €8, 69,
7).
XXVIl
Cyclops
angustus, 5 d57, 68, 69,
16:
annulicornis, 330.
atienuatus, 53, 5d, 68,
75.
bicolor, 54.
bicuspidatus, 70.
ciliatus, 60, 68, 76.
compactus, 62, 68, 77.
cunningtont, 54, 55,
56, 68, 75.
dubius, 63, 68, 6
eminzt, 50, 51, 68, 9,
74.
euacanthus, 59, 62, 68
76.
exiguus, 54, 68, 75
jimbriatus, 61.
hyalinus, 51.
levimargo, 57, 68, 76.
languidus, 329.
leuckarit, 50,
69, 74.
macruroides,
MACTUIUS, =
nanus, 329.
neglectus, 51, 68, 69
74.
oithonoides, 50, 51.
oligarthrus, 61, 68, 77.
pachyc Omes, dd, 68, Ti
phaleratus, 32.
rarispinus, 58, 68, 76.
semiserratus 5b, ~57,
68. 76,
serrudatus, 56, 57, 68
51, 68,
°
336, 321.
teneilus, 52, 68, 75.
varicans, 53, 54, 68,
69, 7a
varias, 59.
vernalis, 329, 3:
viridis, 330, Sail
Cynelurus
jubatus, 117.
Cynictis
selous?, 118.
Cynopithecus, 86.
Cynopterus
brachysoma, 376.
horsfieldi, 376.
titthecheilus, 375.
Cypridopsis
villosa, 330.
Cyrtoma
spuria, 361.
Cysticercus
arionis, 329.
tenié arionis, 329.
INDEX.
TDamaliscus
lunatus, 122.
Danais (Limnas)
chrysippus petilia, 145.
Dasyprocta, 185.
acouchy, 90.
aguti, 90.
azare, 90.
cristata, 516.
prymnolopha, 90.
punctata, 187, 197.
Dasypus
novemeinctus, 90.
sexcinctus. 90.
Dasyurus, 512.
maculatus, 90, 512.
maug@i, 502, 505.
novemeinctus, 505, 507,
509.
setosus, 505, 507, 509.
vellerosus, 505, 506,
507.
Davainea, 328, 331, 332
calva, 351.
cestictllus, 351, 357.
echinohothrida, 355.
globocaudatus, 367,
368.
madagascariensis, 359.
proglottina, 355.
retusa, 566, 368.
ietragona, 300.
urogalli, 351, 352, 354,
ey 360, 361, 362,
366, 367, 368.
ie 366, 368.
Demiegretta
saera, 140.
Dendromus
mesomelas, 120.
Dermestes
Felinus, 151.
Diaptomus
cunningtoni, 37, 68, 73.
galebi, 33, 34.
galeboides, 33, 34, 35,
36, 68, 72.
gracilis, 331.
miztus, 34, 68, 7 =
simplex, 36, 68, 72.
stuhimanni, 35, 37, 38,
72.
Dicerobatis
eregoodoo, 144.
Diceros
bicornis, 124.
Dicotyles, 194, 195, 196.
salvania, 170, 171, 172,
197.
Dicrostonyx
torquaius, 204.
Didelphys, 511, 528.
lanigera, 497.
marsupialis, 497, 498.
pusilla, 497.
virginiana, 90, 498.
Diglossa, 528, 542.
baritula, 527, 5380, 582,
540, 548.
Diodon
hystrix, 406.
Diplax, 254, 284.
Dipodomys, 89.
ambiguus, 90.
spectabilis, 90.
Dipus, 89.
hirtipes, 518.
jaculus, 90.
Dipylidium
caninum, 325.
Distomum
plesiostomum, 366.
Doreatherium, 190.
Drepanis, 543.
Dromezus, 227, 233.
Dynastes
hercules, 414.
Dytiscus, 256.
Echidna, 511, 528.
aculeata, 90.
Hehinorhynchus
stellaris, 366, 368.
Elephantulus
rupestris MYyUrUs,
116.
Elephas, 170, 185, 196.
africanus, 125, 162.
andicus, 90.
maximus, 162.
Emballonura
moniicola, 382, 383.
peninsularis, 383.
Emys
lutaria, 229, 254
Enallagma, 278.
cyathigerum, 255, 266,
267, 269, 272.
Entomyza, 528, 533, 541.
cyanotis, 527, 535, 037,
538, 540, D4].
Epinephelus
striatus, 130.
Equus, 166.
burchellz, 415.
— chapmani, 124.
Foai, 418.
grevyi, 418, 419.
guagga, 418.
— burchelli, 415, 416.
— chapmanni, 416.
wahlbergi, 416.
114,
Equus
zthra, 308, 418.
Erebia, 422.
Ergasiloides, gen. nov.,
63, 67, 69, 71.
brevimanus, (6, 68, 77.
macrodactylus, 65, 68,
Gide
megacheir, 64, 66, 65,
Hie
Ergasilus, 63, 64, 71.
sp., 66, 68, 69.
Prinaceus, 511.
algirus, 510, 512.
europaeus, DLO, 512,
521, 522, 525.
Eriphia
levimana, 159.
Erythromma, 278.
niias, 255, 257, 266,
269, 272.
Esox
lucius, 26.
Endyptula
minor, 226.
Euruppellia
annulipes, 159.
Eurycercus
lamellatus, 330.
Everes, 473.
argiades, 456.
Felis, 497.
carwceal, \\7.
domestica, 494.
javanensis, 385.
leo, 117.
ocreata, 117.
pardus, 117, 206, 384.
— fontanieri, 209.
— leypardus, 205, 206,
207, 209.
— nonopardus, 209.
— parduz, 206, 207,
209.
sp., 367. :
honasi@, 365, 367.
smithi, 349.
urogalli, 367,
Pregata
aquila, 140,
ariel, \A0.
Funisciurus
cepupi, 119.
Fusearia
compar, 364.
refler, 369.
strumosa, 36D.
INDEX,
Galago, 185.
crassicaudatus, BY,
mossambicus, \\4.
senegalensis, 89.
Galeopterus
wariegatus, 388A.
Galictis
harbara, 90.
vittala, BO.
Galidia, 497, 507.
elegans, AT7-496.
Galidictis, 477, 478, 480,
43], 482, 484, 485,
486, 487, 493.
striata, 4177, 483.
Gallinula
chlovopus, 325, 326.
Gallus
gallinaceus, 336.
Gamasus
coleoptratorum, 327.
Gammarus
piler, BB\.
Gastrochata
cybeutes, ANI.
mabille, 411.
Gastrophysa, 281.
Gazella, 160, 197.
rufifrons, \87.
Gecarcoidea
lalandéi, 159.
Gebyra
mutilata, 148.
Gelasimus
sp., 159.
Genetta
ludia, 117.
rubiginosa, \17, 431.
vulgaris, 483, 489.
Geograpsus, 159.
Georhynchus, 521.
capensis, 518.
Georychus
hottentottus, 1.
nimrodi, 122,
Geotomus
PY ymMeUs, 153.
Gerbillus, 521.
egy pius, AT, BVI.
leucogaster, 120.
Giraffa
comelopardalis typica,
309.
capensis, 124.
Glyphidodon
sordidus, 404.
Gly phodes
indica, 149.
Gobius
albbopunctatus, 404.
XxX1X
Goniodes, 313, 315, 316,
318, 319, 320,
heteroceros, 3\\.
tetraonis, 310, BA,
333, 304.
Gonodactylus
chiragra, 160.
Gorgyra
wurde, ANN.
afikpo, ANN, 413.
Graphiurus
griselda, \19.
muninns, NB.
platyops, \N9.
Gryllacris
sp., 157.
rufovaria, 157.
sigmiferda, 157.
Gryllodes
sigillatus, 57.
Gygis
candida, 137, 138, 129,
142,
Hemulon
flavolineatum, 132,
Halicyclops, 62.
Hapale
jacchus, W).
Haplodactylus, 404.
Harpiocephalus
harpia, 331.
lasywrus, 38).
Helictis
orientalis, Bb.
personata, 493, 494.
Helogale
parvila, 118.
Hemerobius
ep., 156.
Hemiclepsis, 19, 24, 29.
Hemigalidia, 477.
Herpestes, 439.
Sulvescens, 482.
mungo, WH.
pulverulentus, 435.
smithi, 494.
Herpetocy pris
tumefata, 30.
Herse
convolvuli, 148.
Hesperia
spi0, 407.
Heterakis
borealis, B6A, 367.
compar, 363, 364, 265,
367.
infleca, 369.
maygnopupila, 366, 567.
pa pillosn, 348, 363, 364,
365, 367.
xXxXX
Heterakis
perspicillum, 363, 365,
366, 367.
vesicularis, 348, 364.
Heteropoda
venatoria, 158.
Hexamitus, 129.
Hippopotamus
amphibius, 124.
Hipposideros
bicolor, 379.
caffer, 115.
diadema, 379.
larvatus, 379.
nobilis, 379.
Hippotragus
equinus, 123.
mger, 123.
Hirundo
gutturalis, 142.
Histiophorus
gladius, 144,
Holaster
planus, 306.
Holocystis
elegans, 292.
Homo
sapiens, 340, 521.
Hyena
crocuta, 118, 495.
striata, 479.
Hydrobius
Juscipes, 261.
Hydrocheerus, 185, 189.
capybara, 516.
Hydromys, 520.
chrysogaster, 516, 517.
Hylobates
leuciscus, 372.
Hymenolepis, 328, 331.
diminuta, 360.
microps, 346, 351, 358,
359, 362, 363, 366,
367, 368.
nana, 360,
tetraonis, 358, 3866,
368
Hypena
strigata, 147.
Hypoleucis
arela, 412.
enantia, 4135.
Hypolymnas
bolina, 144.
— nerina, 145.
misippus, 145.
Hypsiprymnus
sp., 497.
Hyrax, 160, 185, 495.
brucei, 168.
INDEX.
Hyrax
capensis, 162, 163, 165,
166, 167, 168, 169,
196, 5138.
dorsalis, 162, 163, 164,
166, 167, 168, 169,
illeve
Hystrix
sp., 122.
cristata, 516, 522.
javanica, 391,
Ichthyosporidium, 399,
401, 402.
gasterophilum, 400.
phymogenes, 400.
Tctonyx, 507, 524, 525.
capensis, 119, 491, 492,
494, 522,
Iguana, 227.
Ilyophilus
flexibilis, 48, 70, 71.
perplexus, 47, 68, 71,
74.
TIschnura, 277.
elegans, 255, 256, 257,
258, 266, 269, 272,
276, 285.
Ismene
wnicolor, 412.
Tsometrus
maculatus, 158.
Tsoptera, 156.
Ixodes
hexagonus inchoatus,
326.
ricinus, 326.
Juncus
articulatus lampro-
carpus, 332.
effusus conglomeratus,
99
OL.
glaucus, 332.
squarrosus, 332.
Katreus
johnstoni, 407, 408.
Kerivoula
fusca, 382.
hardwicke?, 382.
picta, 381.
Kobus
ellipsiprymnus, 125.
Lacerta, 227.
Lagopus
albus, 311, 319, 321,
364.
alpinus, 319.
mutus, 367, 368.
Lagopus
saliceti, 310, 364.
scoticus, 309, 310, 335,
351, 352, 366, 367,
368.
subalpinus, 364.
tetrix, 310.
Lagostomus
trichodactylus, 510.
516.
Laophonte, 46.
Lariscus
insignis, 389.
— javanus, 389.
Jjalorensis, 389.
peninsulé, 389.
Lasiurus
pearsoni, 381.
Latris, 404.
Leggada
minutoides, 121.
Lemur, 86, 182.
catta, 514, 515.
varius, 87, 88.
Leocyma
sericcata, 146.
tibialis, 146.
Lepidodactylus
lugustris, 143.
Leptis, 352.
Leptocoma, 528.
grayt, 527, 537, 540.
Leptodera
angtostoma, 329.
appendiculata, 329.
Leptodius
sanguineus, 159.
Leptograpsus, 159.
Leptoptilus, 227.
Lepus
caniculus, 340.
nigricollis, 391.
timidus, 340.
euluensis micklemi
122.
Lestes
sponsa, 255.
Leuciscus
erythrophthalmus, 29
Leucocytozoon
lovati, 309.
Leucophcea
surinamensis, 156
157.
Ligula
reptans, 307.
Limax
agrestis, 359.
cinereus, 355.
flavus, 355.
variegatus, 355.
Limenitis, 282.
Liolophus
planisstmus, 159.
Liomera
pubescens, 159.
Lioxantho
punctatus, 159.
Liparis
vulgaris, 400.
Loboptera
sp., 156.
Lophophorus
impeyanus, 320.
Lophophyllum
dumonti, 285.
Loris, 185.
Lutra, 495.
cinerea, 387.
Lycsenopsis
acesina, 428, 452, 433,
476.
aga, 429, 431.
akasa, 458, 460, 476.
albidisca, 446, 447,
471, 476.
alboceruleoides, 446,
476.
albocerulea, 428, 444,
470, 471, 476.
ananga, 419, 429,
4531.
argianus, 419.
argiolus, 419, 420, 421,
423, 428, 437, 443,
444, 445, 456, 468,
476.
— puspargiolus, 446.
— sikhkima, 444.
— victoria, 444, 446,
~ 476.
beretava, 450, 476.
biagt, 451, 456, 457,
458, 476.
binghamt, 428, 476.
bothrinoides, 446, 476.
cajaya, 438, 440, 476.
camene, 428, 456, 458,
460, 461, 476.
cara, 432, 476.
cardia, 451, 452, 458,
456, 462, 476.
carna 470, 476.
catreus, 422, 432, 471,
472, 476.
ceyx, 470, 476.
chennellii, 444, 446,
476.
cinctata, 424, 429, 430,
476.
coalita, 451, 476.
celestina, 444, 476.
INDEX.
Lyczenopsis
corythus, 423, 424, 427,
428, 432, 476.
cossa, 453, 455.
coss@a, 453, 456, 476.
eyanescens, 438, 446,
476.
damme, 4838, 442,
476.
deliciosa, 473.
dilecta, 423, 428, 451,
452, 4538, 454, 455,
458, 462, 476.
dilectissima, 450, 451,
462, 463, 468, 470,
476.
drucei, 458, 459, 476.
duponchellii, 441.
haraldus, 419,
437, 476.
hersilia, 471, 476.
hugelii, 444, 468, 476.
imperatrix, 438, 440,
476.
Jynteana, 444, 446, 447,
450, 476.
kuhni, 438, 476.
ladon, 443, 476.
ladonides, 444, 476.
lambi, 488, 441, 449,
476.
lanka, 428, 456, 457,
436,
476.
levettii, 444, 476.
limbata, 422, 428,
428, 444, 445, 447,
449, 450, 468, 470,
476.
— jynteana, 424, 444.
— placida, 444.
lugra, 422, 458, 476.
lyce, 447, 450, 476.
lyseas, 447, 450, 476.
marginata, 428, 446,
447, 448, 476.
melena, 428, 464, 465,
466, 468, 476.
musina, 422, 423, 427,
498, 432, 458, 476.
nedda, 423, 427, 428,
429, 430, 431, 432,
434, 470, 476.
oreas, 427, 4382, 434,
435, 468, 476.
owgarra, 466, 467, 468,
476
phillippina, 427, 428,
432, 434, 476.
phuste, 432, 476.
placida, 447, 450, 468,
476.
XXXI1
Lycnopsis
placidula, 428, 429,
434, 465, 466, 468,
470, 476.
plauta, 428, 462, 468,
464, 470, 476.
pseudargiolus,
445, 476.
puspa, 423, 427, 434,
437, 438, 439, 440,
441, 442, 443, 444,
446, 476.
puspargiolus, 446, 476.
puspinus, 438, 476.
ripte, 434, 436, 476.
rona, 451, 456, 457,
476.
selma, 423, 458, 461,
462, 476.
semiargus, +19.
445,
shelfordi, 428, 425,
429, 476.
sikkima, 444, 446,
476.
stngalensis, 434, 455,
466, 468, 469, 470,
476.
sonchus,
476.
splendens, 438, 476.
strophis, 447, 450, 468,
476.
tenella, 451, 464, 465,
466, 467, 468, 470,
462, 464,
476.
timorensis, 438, 476.
transpecta, 424, 428,
432, 485, 462, 476.
vardhana, 422, 423,
476.
victoria, 444, 446, 476.
Lycaon
pictus, 119.
Lycosa
sp., 158.
Lygus
sp., 154.
Lyperosomum
corrigia, 366, 368.
Lyrurus
(Letrao) tetrix, 346.
Lysiurus
_wnicinctus, 507, 508.
Macaca
fascicularis, 378.
Macacus, 86.
rhesus, 84.
Macroglossa
passalus, 147.
XXXII INDEX.
Macroglossus Mellivora Mungos
minimus, 376. typicus, 396. Javanicus, 386.
Macropus Meloé, 281. Munia
agilis, 504. Menopon oryztvora, 141.
bennetti, 497, 498. pallescens, 319, 334. Muntiacus
giganteus, 90, 497. pallidum, 318. muntjak, 392.
hagenbecki, 504. persignatum, 312. Murzena
parryt, 503, 504, 505. titan, 316. nebulosa, 403.
robustus, 90. Mephitis, 89, 395, 507. Murina
ruficollis, 90. mephitica, 491, 492, balstont, 381.
rufus, 498. 494. Mus
Madoqua, 173, 181, 182, mesomelas, 90. sp., 391.
186, 189, 197. Merula, 539. auricomis, 12).
phillips, 160, 170, 174, erythropleura, 141. chrysophilus, 120.
183, 188, 190, 191, | Mesoplodon colonus, 121.
192: bidens, 95. coucha, 520.
Madrepora Micraster decumanus, 133, 1854,
centralis, 294. cor-anguinum, 306. 136.
Manatus, 168. Microcebus, 514. Javanus, 391.
Manis, 525. smithii, 515. macleari, 101.
javanica, 392. Microtus microdon, 121.
temmincki, 125. amphibius, 326. musculus, 121,136, 391.
Mantis glareolus, 326. nativitatis, 102.
religiosa, 279, 284. Mimus, 534. neglectus, 390.
siccifolius, 104. Miniopterus nigricauda, 120.
Marasmia blepotis, 382. norvegicus, 391.
venihalis, 148. medius, 382. rattus, 102, 120, 138,
Mazama schreibersi, 116. 134, 186, 390.
rufa, 90. tibialis, 382. Musea
Megaderma Mino domestica, 324.
spasma trifolium, 379. dumonti, 130. Mustela
Megaloperdix Mirabilis erminea, 326.
nigelli, 366. jalapa, 105. Suro, 90.
Megapenthes Moina, 67. vulgaris, 826.
sp., 152. Molytria Mycteroperca
andrewsi, 152. sp., 156. bowersi, 131.
Megisba, 421. Monitor, 234. Mydaus, 386.
malaya, 423, 474, 476. niloticus, 228. Jeoanensis, 387.
Melanoxanthus Monoearya meliceps, 387.
dolosus, 152. centralis, 297, 298, Myopotamus
litura, 152. 304. coypus, 90.
melanocephalus, 152. Monocystis, 309. Myotis
Meles Monodon sp., 381.
meles, 492. monoceros, 200. muricola, 381.
Melia Monophilus (Leuconoe) adversus,
tessellata, 159. ater, 361. 381.
Meliphaga, 545. Montlivaltia, 286. (—) hasselt?, 381.
Melissoblaptes fairfordensis, 291. Myrmecophaga
sp., 148. painswicki, 291. Jubata, 523.
Mellivora, 525. rugosa, 287.
capensis, 396, 397. Moraria.
concisa, 398. brevipes, 330. Nabis
cottont, 894, 397, 398. Moschus, 187, 188, 190. capsiformis, 154.
indica, 397. moschiferus, 185, 185, | Nandinia, 479, 480.
leuconota, 396, 897, 186. binotata, 90, 486, 489.
398. Motella Nannopus, 47.
mellivora, 396. mustela, 400. Nannosciurus
ratel, 119, 396, 397, | Mungos, 385. melanotis, 389.
398. albicauda, 118. Nasilio
signata, 394, 395, 398, cafer, 117. brachyrhynchus, 114,
414, 491, 525. cauui, 117. 116.
Nasua
narica, 90.
rufa, 90, 493.
Nematodum
limacis atra, 329.
Neopithecops, 421.
zalmora, 423, 475, 476.
Nephila
imperatrix, 158.
Nerius
lineolatus, 150.
Nezara
viridula, 153, 154,
Niphanda, 421.
Nirmus, 314, 318,
320.
cameratus, 311, 319,
320, 334.
quadrulatus, 320.
Nisaétus
pennatus, 414.
Nitoera, 45.
Nogodina
bohemant, 154.
Notarthrinus, 422.
binghamt, 424.
corythus, 425.
lambi, 426.
lugra, 426, 427.
musina, 424, 425, 426.
427.
pryert, 423.
vardhana, 424.
Notiomys
cervinus, 519, 520.
Notodromus
monacha, 330.
Notogonia
subtessellata, 149.
Notoryctes
typhlops, 498.
Nycteris
capensis, 115.
Nycticebus, 185.
javanicus, 373.
Nyctinomus
dilatatus, 333.
tenis, 383.
Nyctipithecus
trivirgatus, 89, 90.
Octomitus, 129.
Orypoda, 159.
Odontomachus
hematodes, 149.
Oechalia
consocialis, 158. 154.
(Hsophagostoma, 356.
Ontbophagus
sp., 151.
Proc. Zoou. Soc. 1909
INDEX,
Onychogale
Frenata, 501, 502.
lunata, 501.
Opatrum
sp.. 152.
dubium, 152.
simplex, 152.
Ophiusa
coronata, 146.
melicerta, 146, 147.
Ophyra
chaleogaster, 150.
Oreotragus
oreotragus, 123.
Ornebius
on Wz
Ornithodoros
monbata, 414.
Ornithomyia
avicularis, 321.
Sringillina, 321,
lagopodis, 321, Bye),
334
Ornithorhynchus, 523.
Orthomorpha
coarctata, 158.
Orycteropus
afer, 125.
Osmodes
adon, 407.
Otis
tarda, 366.
| Otocyon
megatotis, 119.
Otumys
trroratus, 520.
— auratus, 120.
Ourebia, 188, 197.
Ovis, 174, 191, 192.
aries, 840.
laticauda, 340.
orientalis, 180.
Pachyura
murina, 3&4.
Pachyzancla
licarsisalis, 142.
stultalis, 149.
Pamphila
tarace, 412.
Pantula
flavescens, 155, 156.
Papilio
lucretia, 409.
nothus, 413,
Papio, 86.
sp., 114.
(Cynocepbalus) hama- |
dryas, 340.
| Paradoxurns, 89.
9,—_No. XXXVII.
Parasmilia,
XXxXlil
Paradoxurus
hermaphroditus, 90,
481, 489, 510.
javanicus, 885.
niger, 90, 481.
285, 286,
290.
centralis, 287, 288,
294, 295, 300, 306,
307.
cylindrica, 287, 301,
303, 305, 306, 307.
jittont, 287, 288, 289,
295, 296, 297, 299,
800, 301, 306, 307.
granulata, 237, 298,
301, 3805, 306, 307.
gravesi, 987, 300, 301,
805, 306, 307.
mantelli, 287, 301,
302, 304, 305, 306,
307.
monilis, 287, 301, 304,
306, 307.
serpentina, 287, 288
293, 306, 307.
Pardaleodes
astrape, 407.
bule, 407, 413.
incerta, 407.
ligora, 407.
makala, 4138.
rutilans, 407.
vibius, 407.
Parnara
arela, 407, 412.
horbonica, 407.
fatuellus, '407.
flavifasciola, 407, 412,
413.
leucophea, 407.
noctula, 407, 411,
413.
palocampta, 407, 411,
413.
statirides, 412.
subochracea, 407.
tarace, 41 2.
Parupeneus
andrewsit, 403.
bifasciatus, 403.
trifasciatus, 403.
Patella
cerulea, 237.
vulgata, 235-253.
Pedetes
cater, 122
Pelagia
phosphora, 81.
Pelodytes
hermaphroditus, 329.
fo ted
oF
XXXIV
Pentalatoma
consocialis, 155.
Perea
fluviatilis, 12, 13.
Perdix
cinerea, 346,
greca, 366.
Periplaneta
americana, 150,
Perisoreus
atricapillus, 1.
Perodicticus
potto, 89, 90.
Peromyscus, 89.
texensis, 90.
Petalia
javanica, 380.
Petaurista
nitida, 387, 388.
Petaurus
taguanoides, 497, 498,
523.
Petrogale
sp., 498.
czanthopus, 498.
Phacocherus, 194, 195.
ethiopicus, 171, 1938.
Phaethon
fudous, 102, 141.
rubricauda, 141.
Phalanger
ursinus, 523.
Phalangista
vulpina, 90, 497.
Phascologale
penicillata, 497.
Phascolomys |
wombat, 497, 498. |
Phasianus
colchicus, 346.
Phenacodus, 170.
Philodieus
Javanus, 150.
Philopterus
cameratus, 319.
Phisis |
listeri, 157.
pectinata, 157.
Pheenicopterus, 234.
antiguorum, 227.
Phyllium
erurifolium, 103-113.
scythe, 105, 107.
siccifolium, 104,
112.
Physocyclus
globosus, 158.
Pilema
octopus, 81.
Pionoeypris
vidua, 330.
INDEX.
Pipistrellus
abramus, 380.
imbricatus, 380.
tralatitius, 380.
Piscicola, 11, 12.
Pison
hospes, 149.
Pithecops, 421.
Plagiolepis
longipes, 149.
Platychelipus, 47.
Platyglossus
hyrtelii, 404.
Plectopterus
gambensis, 227.
Plesiops
melas, 403.
Pleuronectes
jlesus, 401,
Ploceus
hypoxanthus, 141.
Plusia
chalyctes, 147.
Pomatorhinus
erythrogenys, 130.
Poreula
salvania, 160, 170, 171,
172, 197.
Potamocheerus
ethiopicus, 124.
cheropotamus, 124.
Potamocypris
fulva, 330.
Potatia
acuminata, 11.
Precis
villida, 144, 150.
Presbytis, 86.
aygula, 373.
maurus, 372.
mitratus, 373.
pyrrha, 372.
| Procavia
eapensis, 124.
brucei, 125.
Proeyon
cancrivorus, 90.
lotor, 90, 480, 490.
Prodenia
littoralis, 146.
Pronolagus
ruddi randensis, 122.
Prosthemadera, 528.
541.
nove-hollandie; 527.
5
SY), SBS Gs 5
540, 542.
Proteles, 479.
cristatus, 118.
Pseudacrza
lucretia, 409.
|
| Python
Pseudochirus
peregrinus, 501.
Pseudolfersia
spinifera, 150.
| Pseudoscarus
sp., 144.
Pseudozius
caystrus, 159.
Psilopus, 150.
Pterois
volitans, 404.
Pteropus, 524.
natalis, 102.
vampyres, 373, 375.
Pudna
humilis, 186.
Puffinus
brevicaudus, 142.
sphenurus. 142.
Pulchriphyliuin,
104.
Pulex
serraticeps, 325.
Pycnosoma
jlaviceps, 150.
Pyralis
“nanthotalis, 148.
Pyrrhosoma, 277.
nymphula, 257,
269, 272
ale.
108,
266.
sebe, 204.
| Raja
maculata, 204.
_ Rallus
philippensis, 137
Raphicerus
campestris. 123.
sharpei, 123.
Remigia
Srugalis, 147.
Remipes
testudinarius, 159.
Rhamphastos
ariel, 226.
Rhaphicerns, 188, 197.
Rhea, 227.
Rhinia
testacea, 150.
Rhinoceros
bicornis. 198. 199, 2O0.
holmwaodi, 199.
oswelli, 199.
simus, 198, 199, 200.
cottoni, 198, 200.’
Rhinolophus
acuminaius. 377.
affinis, 377.
augur zaimbesiensis,
lo.
Rhinolophus
borneensis, 376, 377.
canuti, 377, 378.
creaghi, 378, 379.
darlingi, 115.
empusa, 113, 115.
geminus, 377.
hildebrandti, 115.
luctus, 377.
minor, 378.
pusillus, 377, 378.
stheno, 377.
trifoliatus, 377.
Rhopalocampta
bixe, 407.
hanno, 407.
unicolor, 407, 412.
Rhopica
binotata, 153.
honesta, 153.
Ricania
bohemani, 154.
Rousettus
amplexicaudatus, 374.
leschenaulti, 374.
minor, 375.
shortridget, 374, 375.
Saccostomus
hilde, 121.
Salarias
anomalus, 406.
caudofasciatus, 405,
406.
hasseltii, 405.
marmoratus, 405.
melanosoma, 406.
natalis, 405, 406,
unicolor, 405.
Salvelinus
graye, 98.
jmaxillaris, 98.
Sarangesa
honvieri, 407.
hrigidella, 407.
perpa upera, 407,
Sarcophaga
sp., 150.
Scarus
sp., 144.
Scatophaga
sp., d01.
sgualida, 362.
stercoraria, 325, 324,
325, 334, 361.
Schizopera, 39, 49, 68, 69.
consimilis, 42, 68, G9,
loa
fo
jimbriata, 45, 68, 69, 74.
inopinata, 40, 41, 42,
68, 73.
INDEX.
Schizopera
longicauda, 40, 41, 42,
ae |
minuticornis, £3, 44, |
68, 73. |
scalaris, 46, 68, 74.
spinulosa, 44, 45, 68,
TA.
ungulata, 43, 68, 73.
validior, 41, 42, 493, |
68, 73.
Schizophrys
aspera, 159.
Sciuropterus
(Hylopetes) sagitta,
387.
Sciurus, 525, 526.
andrewst, 388.
aurantiacus, 388.
everetti, 388.
genibarbis, 387.
lepidus, 387.
maximus, 518.
nigro-vittatus, 388.
notatus, 388, 389. |
prevosti, 496, 517, 518. |
sagitta, 387, 388.
spadiceus, 388.
vulgaris, 387. |
Scolopax
gallinago, 142.
Scotophilus
kuhli, 381.
migrita, 116.
temminckii, 380, 381.
Scutellina
patella, 309.
Semnopithecus
maurus, 3872.
mitratus, 373.
pyrrhus, 372.
Sessinia
sp., 152. |
andrewsi, 152.
Sialia, 53-4.
Siderastraa, 292.
Simia
aygula, 373.
Simosa
vitula, 330.
Smeringopus
elongatus, 158.
Sorex
vulgaris, 326.
Spheerodon
heterodon, 403.
Spheniscus
demersus, 226, 227.
Sphenodon, 227.
Sphinx
ligustri, 271.
KZXAV
Spirochzeta
duttoni, 4\4.
gallinarum, 327.
Spodoptera
mauritia, 146.
Stegomyia, 151.
Sterna
fuliginosa, 137.
Stizus
sp., 149.
reversus, 149.
Strepsiceros
strepsiceros, 124.
Strongylus
anseris, 340.
contortus, 340.
crisptuus, 340.
douglassti, 339.
gracilis, 335, 340.
hastatus, 366, 367.
nodularis, 340.
papillatus, 366, 367.
perqracilis, 335, 336,
340.
quadriradiatus, 330,
336, 340.
strigosus, 340.
tenwis, 336, 340.
Struthio, 227.
Sula
abbotti, 140.
piscatrix, 141.
sula, 140.
Suricata
tetradactyla, 480, 482,
487, 488, 522.
Sus, 170.
cristatus, 171, 195, 195.
salvania, 196, 197.
verrucosus, 391.
vittatus, 392.
Sympetrum
striolatum, 254, 257,
279:
Syngamus
trachealis, 344, 567.
Teena
bonasie, 365, 368.
botriopliti, 365.
calva, B51, 352.
cesticillus, 357.
echinata, 864, 368.
microps, 308.
urogalli, 301.
Vaphozous
fulvidus, 385.
longimanus, 385.
melanopogon, 383.
saccolaimus, 383.
theobald?, 383.
XXxvi
Tapirus, 160.
americanus, 308.
indicus, 161, 196, 418.
malayanus, 196.
Tarucus
nara, 473.
Tatera
lobenaule, 120.
Tatusia
kappleri, 509, 510,
B25:
peda, 506, 509.
Taurotragus
oryx, 124, 414.
Termites, 281.
Testudo, 228, 233.
greca, 229.
Tetrao
bonasia, 365.
tetra,
567, 368.
urogallius,
365, 367, 368.
Tetrodon
sp.. 144.
patoca, 144.
Thalassochelys
caretia,. 239.
Theeosmilia
obtusa, 291.
(Choriastreea)
291.
Thersites, 63.
Thrixion, 281.
halhidayanum, 284.
Thryonomys
surnderianus, 122.
Thylacinus
cyanocephalus, 444.
Tinea
vulgaris, 28.
Tragelaphus
scriptus, 189.
Tragulus, 181, 189, 190,
191, 197, 524, 525.
focalinus, 392.
meminna, 514.
stanleyanus, 182,
184, 514.
Tramea
rosenbergi?, 155, 156.
Trichocephalus
dispar, 336, 348.
trichiurus, 347, 548.
Trichoglossus
cyanogrammus, 130.
Trichosoma
caudinflatum, 345.
gallinum, 349.
320, 365, 366,
320, 346,
re ‘asa y
83,
INDEX.
Trichosoma
longicolle, 341, 344,
545, 346, 348, 349,
350, 868, 365, 367.
retusum, 346,
Trichostrongylus
capricola, 340.
extenuatus, 340.
instabilis, 340.
nodularis, 336, 340.
pergracilis, 335, 356,
3de7, 340, 341, 342,
345, 344, 348, 349,
350, 367.
probolurus, 340.
quadriradiatus, 340.
retorieformis, 340, 342.
subtilis, 340.
tenuis, 336, 340.
vibrinus, 340.
(Strongylus) nodularis,
300.
(—) tenuts, 335.
Trichosurus
fuliginosus, 499, 500,
505.
vulpecula, 498, 499,
501, 502, 523.
Trigoniulus
sp., 158.
Tripterygium
atrogulare, 404.
Troehilus, 530, 532, 533.
Trochoideus
desjardinsi, 151.
Trypanoplasma, 3, 10.
abramidis, 8, 29, 30.
borrelt, 4, 9, 29, 30.
cyprint, 4.
gurneyorum, 5, 23, 24,
25,26, 27, 29; 30.
Keysselitzi, 23, 26, 27,
28, 29, 30:
Terre
abramis, 24, 30.
carassit, 23.
eberthi, 309.
granulosum. 8, 14,
19, 25, 29, 30.
lewisi, 4, 8, 10, 18.
magna, 7, 8, 19,
remaki, 7, 8,
24, 30.
tince, 23, 24, 27, 30.
Tubifex, 7.
Tupaia
javanica, 584.
Tupinambis
teguexin, 227.
Turdus, 370.
merula, 325.
Tylenchus
tritic?, 348.
Tylonycteris
pachypus, 380.
Typhlops
braminus, 143.
Tyroglyphus
farine, 327.
Urocissa
occipitalis, 130.
Ursus
arctos, 1.
syriacus, 493, 494.
Utethesa
pulchelloides, 145.
Vanellus
vanellus, 825.
Vanessa
kershawi. 145.
Varanus, 170, 228.
Vespertilio
Serrum-equinum minor,
92-Q
od.
minutus, 115.
Viverra
etvetta, 117, 481, 484.
489, 495.
megasprla, 90.
Viverricula
indica, 90.
rasse, 389.
Vulpes
leucopus, 130.
Zaphrentis
edwardsiana, 285.
vermicularis, 285.
Zapus, 521, 526.
hudsonianus, 51 7,519.
Zinckernia
Sascialis, 148.
Zizera
alsus, 422.
maha, 422.
minimus, 422.
Zosterops
natalis, 141.
Printed by Tayror and Frascis, Red Lion Court, Fleet Street.
—
Se
No. 64.
ABSTRACT OF THE PROCEEDINGS
ZOOLOGICAL SOCIETY OF LONDON.
January 12th, 1909.
Prof. J. Rose Braprorp, M.D., D.Sc., F.B.S., Vice-President,
in the Chair.
The Secretary read a Report on the additions that had been
made to the Society's Menagerie during the month of December
1908.
Dr. H. G. Puumuer, F.LS., F.Z.8., Pathologist to the Society,
exhibited the Intestinal Tracts of two Snakes that had recently
died in the Society's Gardens and called attention to the condition
of inflammation present in them.
Prof. E. A. Miscuts, M.A., V.P.ZS., read a paper entitled
“ Observations on the Flagellates Parasitic in the Blood of Fresh-
water Fishes,” in which five species of Trypanosoma and four
species (two new) of Trypanoplasma, from fishes of the Norfolk
Broads, were described in detail. Particular attention was paid
to the minute structure of the parasites, and it was shown that
it is possible to give a uniform description for the nuclear
apparatus of both Trypanosoma and Trypanoplasma.
Dr. W. A. Cussisetos, F.Z.S., read a paper by Prof. G. O. Sazs,
C.M.ZS., entitled “ Zoological Results of the Third Tanganyika
Expedition, 1904-1905.— Report on the Copepoda.”
* This Abstract is published by the Society at 3 Hanover Square, London,
W., on the Tuesday following the date of Meeting to which it refere. It will
be issued, free of extra charge, to all Fellows who subscribe tothe Publications,
along with the ‘Proceedings’; but it may be obtained on the day of publication
at the price of Sixpence, or, if desired, sent post-free for the eum of Siz
Shillings per annum, payable in advance,
2
Tn a paper communicated by Prof. T. W. Bripex, D.Sc., F.B.S.,
F.Z.S., and entitled “‘On the Gonadial Grooves of a Medusa,
Aurelia aurita,” the author, Mr. T. Goopry, B.Sc., Zoological
Laboratory, University of Birmingham, dealt with investigations
which confirmed his earlier suggestion that the gonadial grooves,
which lie in the interradial axes between the central gastric
cavity and the gastric pouches, have a sexual function. From
sectionised material, drawings had been obtained of spermatozoa
and eggs lying within the limits of the gonadial grooves, thus
proving that the latter are functional gonoducts.
The SrecreTary, on behalf of Mr. Artaur Erwin Brown,
D.Sc., O.M.Z.S., Secretary of the Zoological Society of Phila-
delphia, read a paper entitled “‘ The Tuberculin Test in Monkeys ;
with Notes on the Temperature of Mammals.” The paper
described the methods and results of experiments which have
recently been carried out at the Zoological Gardens of Phila-
delphia with the view of suppressing tuberculosis in monkeys.
Mr. F. E. Bepparp, F.R.S., F.Z.S., presented a paper by Prof.
R. Couterr, F.M.Z.8., “On Balena glacialis and its Capture in
recent Years in the North Atlantic by Norwegian Whalers.”
The next Meeting of the Society for Scientific Business will
be held on Tuesday, the 2nd February, 1909, at half-past Eight
o'clock p.m., when the following communications will be
made :—
1. Dr. C. W. Anprews, F.RB.S., F.Z.8S.—Notes on the Fauna
of Christmas Island, illustrated by lantern-slides.
2. Dr. H. G. Prruuer, F.L.S., F.Z.5.—Report on the Patho
logical Observations at the Society’s Gardens during 1908.
3. H. 8. Lutes.—Preliminary Account of the Life-history ot
the Leaf-Insect, Phylliwm crurifolium Serv.
4, . CO. Cuuss, F.Z.8.—The Mammals of Matabeleland
3
The following communications have been received :—
1. F. Woop-Jonzs, B.Sc., F.Z.8.—The Fauna of the Cocos-
Keeling Atoll.
2. Grovuse-Diszase Commission Reports :
(c) A. E. Sarrrey, M.A., D.Sc., F.R.S., F.Z.8.—The
Ectoparasites of the Grouse.
(6) A. E. Surrey, M.A., DSc, F.RS., F.Z.8.—The
Thread-Worms (Nematoda) of the Red Grouse (Tetrao
scovicus).
(c) A. BE, Suterey, M.A., D.Sc., F.B.S., F.Z.S.—The Tape-
Worms (Cestoda) of the Grouse. Appendix: Parasites of
Birds allied to the Grouse,
Communications intended for the Scientific Meetings of the
ZooLOGICAL Society oF Lonpon should be addressed to
P. CHALMERS MITCHELL,
Secretory.
3 Hanover Square, Loxpox, W.
January 19th, 1909,
* Say SWAN a fee Vera Uys vi
ue sta a bn. Dy A
See me Or tata eed Uses | Via
1 Ns A
LO, ROAD, Rae CO et eee
Pi) dee
-
¥ a
y a ret a oe ‘i hy
r ; * ‘
Steins eon
No. 65.
ABSTRACT OF THE PROCEEDINGS
OF THE
ZOOLOGICAL SOCIETY OF LONDON.*
February 2nd, 1909.
Freverick Giuurtr, Esq., Vice-President, in the Chair.
Mr. C. Tare Reean, M.A., F.Z.S., exhibited specimens of the
Char of Lough Melvin (Salvelinus grayi, Giinth.) and of the
Char froma little loch under Ben Hope, Sutherlandshire, recently
described by him under the name Salvelinus mawillaris. He
pointed out the differences between the two forms, and called
attention to the interest attaching to the study of this too much
neglected group of British freshwater fishes.
Mr. R. E. Hotprne exhibited several skulls and photographs
of the St. Kilda or Hebridean Four-horned Sheep, and remarked
upon the character of the horns of this curious form, pointing
out some differences between them and those of the South-African
and other Sheep having four horns.
Mr. C. Davies SHerBorn, F.Z.S., on behalf of Mr. Matcoum
MAcLAREN, read an account of a fight between a Whale and a
Swordfish observed in the Hauraki Gulf, New Zealand.
Dr. C. W. Anprews, F.R.S., F.Z.S., gave an account, illus-
trated by lantern-slides, of his visits to Christmas Island, calling
attention to the differences in the fauna associated with influx of
population.
* This Abstract is published by the Society at 3 Hanover Square, London,
W., on the Tuesday following the date of Meeting to which it refers. It will
be issued, free of extra charge, to all Fellows who subscribe to the Publications,
along with the ‘ Proceedings’; but it may be obtained on the day of publication
at the price of Siapence, or, if desired, sent post-free for the sum of Sic
Shillings per annum, payable in advance.
6
Mr. H. 8. Lereu, Honorary Research Fellow in the University
of Manchester, presented a paper, communicated by Prof. Hickson,
F.RS., F.Z.8., entitled “ Preliminary Account of the Life-history
of the Leaf-Insect, Phylliwm crurifoliwm Serville,” and made the
following remarks :—
“The Leaf-Insects occur in the tropical regions of the Old
World and seem partial to insular life. The eggs, which
resemble the seeds of certain plants to a remarkable degree,
require to be kept in a constantly warm and moist atmosphere
to enable them to hatch; they hatch very irregularly, and the
period of incubation often extends over three or four months.
When young the larve are active as compared with older indi-
viduals. The metamorphosis is incomplete, and the adult form
is attained by a gradual increase in size; fully-developed tegmina
and wings only appear in the adult condition. The adult females
are large and leaf-like in appearance, but the males are much
smaller and not foliaceous.”
Mr. E. C. Cuuss, F.Z.S., Acting Curator of the Rhodesia
Museum, communicated a paper on “'The Mammals of Matabele-
land.”
Dr. H. G. Purmer, F.LS., F.Z.8., Pathologist to the Society,
reported on the Pathological Observations at the Society’s
Gardens during 1908, and illustrated his remarks with a series
of lantern-slides.
The next Meeting of the Society for Scientific Business will
be held on Tuesday, the 16th February, 1909, at half-past Hight
oclock P.mM., when the followmg communications will be
made :—
1. F. Woon-Jonges, B.Sc., F.Z.8.—The Fauna of the Cocos-
Keeling Atoll.
2. F. E. Bepparp, M.A., F.R.S8., F.Z.8.—Contributions to the
Anatomy of certain Ungulata, including Zapirus, Hyrax, and
Antilocapra.
3. Prof. Dr. E. L. Trovuressarr, C.M.Z.8.—Le Rhinoceros
Blane du Soudan.
7
The following communications have been received :—
1. Grovsz-DisEASE Commission REPORTS :
(a) A. E. Surprey, M.A., DSc. F.R.S. F.Z.S.—The
Ectoparasites of the Grouse.
(6) A. EH. Surptey, M.A., D.Sc, F.R.S., P.Z.S.—The
Thread-Worms (Nematoda) of the Red Grouse (Yetrao
scoticus).
(c) A. E. Surpney, M.A., D.Sc. F.R.S., -F.Z.S.—The
Tape-Worm (Cestoda) of the Grouse. Appendix : Parasites
of Birds allied to the Grouse.
2. Frank BAurour-Browne, M.A., F.R.S.E., F.Z.8.
The Life-History of the Agrionid Dragonfly.
Communications intended for the Scientific Meetings of the
ZooLocicaL Society oF Lonvon should be addressed to
P, CHALMERS MITCHELL,
Secretary.
3 Hanover Square, Lonpon, W.
February 9th, 1909.
No. 66.
ABSTRACT OF THE PROCEEDINGS
OF THE
ZOOLOGICAL SOCIETY OF LONDON.
February 16th, 1909.
Freperick Gittert, Esq., Vice-President, in the Chair.
The Sucrerary read a report on the Additions made to the
Society’s Menagerie during the month of January 1909.
Mr, C. Tare Recan, M.A., F.Z.S., exhibited sketches illus-
trating colour-changes in sone Fishes from the Bermudas in the
New York Aquarium.
Mr. E. G. B. Meape-Watpo, F.Z.S., read extracts from a
letter he had_received from Dr. Einar Lonnberg, C.M.Z.S., on
the hunting of the Sea-Elephant on South Georgia, and called
attention to the necessity of steps being taken to prevent its
extermination.
Dr. F. Woop-Jonus, B.Sc., F Z.S., presented a communication
on the “ Fauna of the Cocos-Keeling Atoll,” and illustrated his
remarks with lantern-slides. The work was based on collections
made by the author during a stay of fifteen months in 1905 and.
1906, and in the case of most orders was believed to be fairly
complete.
Dr. Wood-Jones had been assisted by various authorities: the
* This Abstract is published by the Society at 8 Hanover Square, London,
W., on the Tuesday following the date of Meeting to which it refers. It will
be issued, tree of extra charge, to all Fellows who subscribe to the Publications,
along with the ‘ Proceedings’; but it may be obtained on the day of publication
at the price of Siapence, or, if desired, sent post-free for the sum of Sz
Shillings per annum, payable in advance.
10
species of Reptilia were determined by Mr. G. A. Boulenger,
F.R.S., V.P.Z.8.; of Lepidoptera Rhopalocera by Mr. F. A.
Heron; of Lepidoptera Heterocera by Sir G. F. Hampson, Bt.,
F.Z.8.; of Hymenoptera by the late Col. C. T. Bingham, F.Z.S. ;
of Diptera by Mr. E. KH. Austen, F.Z.S.; of Coleoptera by
Messrs. C. O. Waterhouse, P.E.S., C. J. Gahan, F.E.S., and
G. J. Arrow, F.E.S.; of Rhynchota by Mr. W. L. Distant, F.E.S. ;
of Neuroptera and Orthoptera by Mr. W. F. Kirby, F.L.S., F.E.S.;
of Arachnida and Myriopoda by Mr. A. S. Hirst, F.Z.S.; and of
Crustacea by Dr. W. T. Calman, F.Z.8.
A paper was communicated by Mr. F. E. Brepparp, M.A.,
F.R.S., F.Z.S., Prosector to the Society, on ‘The Anatomy of
certain Ungulata, including Vapirus, Hyrax, and Antilocapra.”
Dr. E. L. Trovessart, C.M.Z.S., presented a paper entitled
“ Le Rhinocéros Blane du Soudan (Ahinoceros simus cotiont).”
The next Meeting of the Society for Scientific Business will
be held on Tuesday, the 2nd March, 1909, at half-past Hight
o'clock p.M., when the following communications will be
made :—
1. Miss Marcaret Pootz.—The Development of the Sub-
divisions of the Pleuro-peritoneal Cavity in Birds, illustrated by
lantern-slides.
2. E.S. Russett, M.A.—The Growth of the Shell of Patella
vulgata, L.
3. Frank Batrour-Browne, M.A., F.R.S.E., F.Z.8.—The
Life-history of the Agrionid Dragonfly.
4. W. D. Lane, M.A., F.Z.8.—Growth-stages in the British
Species of the Coral Genus Parasmilia.
11
The following communications have been received :—
1. Grouse-DisEasE CommirrEE Reports :
(a) A. E. Surprey, M.A., D.Sc., F.R.S., F.Z.S.—The
Ectoparasites of the Grouse.
(6) A. EH. Suiprey, M.A., D.Sc, F.R.S., F.Z.S.—The
Thread-Worms (Nematoda) of the Red Grouse (TZetrao
scoticus).
(c) A. KE. Surprey, M.A., D.Sc., F.R.S., F.Z.8.—The Tape-
Worms (Cestoda) of the Grouse. Appendix: Parasites of
Birds allied to the Grouse.
2. W. P. Pycrart, F.Z.S., A.L.S.—On a Fossil Bird from the
Lower Pliocene.
Communications intended for the Scientific Meetings of the
ZOOLOGICAL Society oF Lonvon should be addressed to
P. CHALMERS MITCHELL,
Secretary.
3 HANovER Square, Lonpon, W.
february 23rd, 1909,
Aber
CRS ee AEE AR eS a
Beg Fa.
No. 67.
ABSTRACT OF THE PROCEEDINGS
OF THE
ZOOLOGICAL SOCIETY OF LONDON.
March 2nd, 1909.
G. A. Bounencer, Esq., F.R.S., Vice-President, in the Chair.
Mr. R. H. Burne, F.Z.8., exhibited specimens of elastic
mechanisms in Fishes and a Snake which had been prepared for
the Museum of the Royal College of Surgeons.
Dr. R. F. Scuarrr, BSc., F.Z.S., exhibited a number of
Reindeer bones and antlers, obtained from Ivish caves, which
displayed marks showing that they had been gnawed by different
kinds of animals, probably in some cases by Rodents.
Mr. R. I. Pocock, F.L.S., F.Z.S., exhibited the skulls of some
Leopards, and called attention to the differences in skulls from
Africa and India and to the evidence afforded by them as to the
existence of a small and large type of leopard in Africa.
Miss Marearer Poote read a paper on “The Development of
the Subdivisions of the Pleuro-peritoneal Cavity in Birds,” com-
municated by Prof. G. C. Bourne, D.Sc., F.Z.8., and illustrated
her remarks with lantern-slides.
A paper was received from Mr. EK. 8. Russrtt, M.A., entitled
“The Growth of the Shell of Patella vulgata L.,” communicated
* This Abstract is published by the Society at 3 Hanover Square, London,
W., on the Tuesday following the date of Meeting to which it refers. It will
be issued, free of extra charge, to all Fellows who subscribe to the Publications,
along with the ‘ Proceedings’; but it may be obtained on the day of publication
at the price of Stxpence, or, if desired, sent post-free for the sum of Six
Shillings per annum, payable in advance,
14
by Prof. J. AnrHuR THomson, F.Z.8., of which the following is a
summary :—
The breeding-season of this limpet extends from July to
January. Sexual maturity is reached at a length of 20-25 mm.
An average size for a limpet of the last season’s brood in January
or February is 10 mm.; at the end of the first year it may be
29 mm. Probable sizes at the end of the second and subsequent
years are 38, 44, 48, 53mm. Shells over 50mm. may be
considerably more than five years old.
Sexual maturity is reached in the first year and when the
limpet is only half-grown. The rate of growth decreases with
age and maturity, and is slower during the colder months of the
year. Considerable changes take place in the ratios of the shells’
dimensions during growth, being probably in large part the
expression of “laws of growth” and not due to natural
selection.
Mr. Frank Batrour-Brownge, M.A., F.R.S.E., F.Z.8., presented
a paper on “ The Life-history of the Agrionid Dragonfly.”
Mr. C. Davies Surrporn, F.Z.8., communicated a paper by
Mr. W. D. Lane, M.A., F.Z.S., entitled “‘ Growth-stages in the
British Species of the Coral Genus Parasmilia.”
The next Meeting of the Society for Scientific Business will
be held on Tuesday, the 16th March, 1909, at half-past Hight
o'clock p.M., when the following communications will be
mace :—
1. Grovusse-DIsEASE CoMMITIEE REPORTS:
(a) A. BE. Suieuny, MA.) D.Se.,) FIRS.) B28: The
Ectoparasites of the Grouse.
(6) A, E. Suipiny, M.A,, D.Sc., F.R.S., F.Z.S.—The
Thread-Worms (Nematoda) of the Red Grouse (Teérao
scoticus).
(c) A. KE, Surprey, M.A., D.Sc., F.B.S., F.Z.8.—The Tape-
Worms (Cestoda) of the Grouse. Appendix: Parasites of
Birds allied to the Grouse.
2. W. P. Pycrart, F.Z.S., A.L.S.—On a Fossil Bird from the
Lower Pliocene.
15
3. OuprietD THomas, F.R.S., F.Z.S., and R. C. Wrouacuton,
F.Z.S.—On a Collection of Mammals from Western Java,
presented to the National Museum by Mr. W. H. Balston.
The following communications have been received :—
1. Miss Murret Rosertson.—Notes on an Ichthyosporidian
causing a Fatal Disease in Sea-Trout.
2. C. Tare Recan, M.A., F.Z.8.—A Collection of Fishes made
by Dr. C. W. Andrews, F.R.S., F.Z.S., at Christmas Island.
Communications intended for the Scientific Meetings of the
ZOOLOGICAL Society oF Lonpon should be addressed to
P, CHALMERS MITCHELL,
Secretary.
3 Hanover Square, Lonpon, W.
March 9th, 1909.
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No. 68.
ABSTRACT OF THE PROCEEDINGS
OF THE
ZOOLOGICAL SOCIETY OF LONDON.*
March 16th, 1909.
FREDERICK GILLETT, Esq., Vice-President, in the Chair.
The SEcrEerary read a report on the Additions made to the
Society’s Menagerie during the month of February 1909.
The Secrerary exhibited, on behalf of Mr. E. C. Couss, F.Z.S.,
the skins and skulls of two foetal lions which had been presented
to the Rhodesia Museum, Bulawayo, by Mr. A. Giese. They had
been taken from a lioness which Mr. Giese shot last November at
Deka, about 50 miles south of the Victoria Falls.
These foetuses showed very little pattern compared to that of
newly born cubs; the black markings in each example of the
former consisting only of a fairly well-defined median dorsal line,
a mottling on the head, some distinct spots on the outer sides of
the limbs, and a suspicion of rings on the tip of the tail.
The Secrerary exhibited a photograph of a young American
Tapir, and called attention to the remarkable resemblance between
that and the young Malayan Tapir, a photograph of which was
reproduced in the Society’s ‘ Proceedings,’ 1908, p. 786. The
longitudinal light stripes on the flanks of the body, the spots on
the legs, and the white tips to the ears were present in both.
The Secretary called attention to an interesting paper by
C. ONELLI in the ‘ Revista del Jardin Zoologico de Buenos Aires,’
1908, p. 207, in which the author called attention to a general
correspondence between the number of vertebra and the number
of stripes or rows of spots in many mammals.
* This Abstract is published by the Society at 8 Hanover Square, London,
W., on the Tuesday following the date of Meeting to which it refers. It will
be issued, free of extra charge, to all Fellows who subscribe to the Publications,
along with the ‘ Proceedings’ ; but it may be obtained on the day of publication
at the price of Stxpence, or, if desired, sent post-free for the sum of Str
Shillings per annum, payable in advance.
18
The Secretary exhibited a photograph of a small herd of
Mountain Zebras (Hquus zebra) in the possession of a dealer at
Port Elizabeth, South Africa.
The Srcrerary also exhibited a photograph of a female Giraffe
captured in the West Soudan, east of Timbuctoo, and showing a
general resemblance with Giraffa camelopardalis typica.
Dr. F. A. Barusr, M.A., F.Z.8., exhibited a fossil Echinoid,
Seutellina patella, from the Hocene? (Barwonian) of Nelson,
Glenelg River, Victoria, Australia, showing a marsupium for the
young, as described by T. 8. Hall (Proc. Roy. Soc. Victoria, n. s.
xx. p. 140, 1908).
Dr. A. E. Sarpiry, M.A., F.R.S., F.Z.S., communicated the
following Reports of the Grouse Disease Committee :—(a) The
Kctoparasites of the Grouse ; (b) The Thread- Worms (Nematoda)
of the Red Grouse (Zetrao scoticus); (c) The Tape-Worms
(Cestoda) of the Grouse.—Appendix: Parasites of Birds allied
to the Grouse. He gave a general description of the work of the
Committee, and explained the results of the examination of the
parasites of the Grouse, exhibiting drawings and specimens to
illustrate his remarks.
My. W. P. Pycrart, F.Z.8., A.L.S., presented an account of the
fossilized remains of a small Passerine Bird, from the Lower
Pliocene of Gabbro, near Leghorn, which most nearly resembled
those of the living species known as Berthelot’s Pipit (Anthus
bertheloti).
A paper was read by Messrs. OLDriELD Tuomas, F.R.S., F.Z.S.,
and R. C. Wrovueuron, F.Z.S., “On a Collection of Mammals
from Western Java, presented to the National Museum by
Mr. W. E. Balston.”
The Island of Java had been almost entirely neglected during
the last 60 years, while it had been one of the most prolific sources
of early described species, and in consequence workers had been
much embarrassed for want of modern specimens representing
these early species for comparison with their allies elsewhere.
Now, thanks to the generosity of Mr. Balston, a very fine collec-
tion had been made in the island by Mr. G. C. Shortridge, and
presented to the National Museum. It consisted in all of over
1500 specimens, belonging to 74 species, of which six were new.
Of these latter, besides the previously described dlurina
balstont, the following were most worthy of mention :—
RHINOLOPHUS CANUTI, Sp. 1.
Allied &. creaghi, but with a low connecting process between
the sella and the posterior lancet, on which latter there is a
prominent tuft of hair.
Forearm 50mm, Type. B.M. No. 9.1.5.183.
18)
RovUSsETTUS SHORTRIDGEI, sp. n.
Allied to 2. leschenaulti, but considerably larger and with much
larger teeth.
Forearm 94 mm.; greatest length of skull 42 mm. Type.
INCH CEO
LARIA INSIGNIS JAVANA, subsp. n.
Larger, darker in colour and with larger teeth than true insignis.
Hind foot 47 mm.; greatest length of skull 53 mm. Type.
Noe RED lS,
CHIROPODOMYS ANNA, Sp. n.
Near C. gliroides, but with smaller skull and teeth.
Skull length 24 mm.; molar series 3°83 mm. Type. No.
Oe aoe
The next Meeting of the Society for Scientific Business will
be held on Tuesday, the 6th April, 1909, at half-past Hight
o'clock P.mM., when the following communications will be
made :—
1. Miss Murret Ropertson.—Notes on an Ichthyosporidian
causing a Fatal Disease in Sea-Trout.
2. C. Tare Reean, M.A., F.Z.S.—A Collection of Fishes made
by Dr. C. W. Andrews, F.R.S., F.Z.8., at Christmas Island.
3. R. I. Pocock, F.L.S., F.Z.8.—Description of anew Form of
Ratel (J/ellivora) from Sierra Leone, with Notes upon the described
African Forms of this Genus.
4, Hamruron H, Druce, F.LS., F.Z.8.—On some New and
Little-known Hesperide from Tropical West Africa,
The following communications have been received :—
1, T. AugERNon CuHapman, M.D., F.Z.S., F.E.S—A Review of
the Species of Celastrina (Tutt), (Lycenopsis, Felder), (Cyaniris,
auct.), on examination of the male ancillary appendages.
2. F. E. Bepparp, M.A., F.R.S., F.Z.S.—On some Points in
the Structure of Galidia elegans, and on the Postcaval Vein in
Carnivora.
Communications intended for the Scientific Meetings of the
ZOOLOGICAL Society oF Lonpon should be addressed to
P. CHALMERS MITCHELL,
Secretary.
3 ELANOVER SQuarRE, Lonpon, W.
March 23rd, 1909,
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No. 69.
ABSTRACT OF THE PROCEEDINGS
OF THE
ZOOLOGICAL SOCIETY OF LONDON,*
April 6th, 1909.
FREDERICK GILLETT, Esq., Vice-President, in the Chair.
The Secretary exhibited, on behalf of Mr. Grorcre JEnnison,
some fertilized eggs from a pair of Seba Pythons in the Belle Vue
Zoological Gardens, Manchester.
Dr. R. T. Lerper, F.Z.S., exhibited a greatly distorted Elephant’s
tusk from the Congo, and a malformed canine tooth of a Hip-
popotamus from Uganda, the conditions most probably originating
from mechanical injury.
Mr. HK. T. Newron, F.R.S., F.Z.8., exhibited a metatarsal bone
of an Ox, showing in a remarkable manner the marks of gnawing
by rodents, possibly Squirrels, Rats, or Mice, which he had found
in the woods near Cromer.
Mr. R. I. Pocock, F.LS., F.Z.S., communicated a paper
entitled ‘* Description of a new Form of Ratel (Mellivora) from
Sierra Leone, with Notes upon the described African Forms of
this Genus.”
Miss Murret Ropertson read a paper, communicated by
Prof. E. A. Mincuin, M.A., V.P.Z.S., ““On an Ichthyosporidian
causing a Fatal Disease in Sea-Trout.”
Mr. C. Tare Recan, M.A., F.Z.S., read a paper on a small
series of fishes from Christmas Island, collected by Dr. C. W.
Andrews, F.R.8., F.Z.8. Seven new species were described, com-
prising five Blennies, a Pampeneus, anda Cirrhites. In connection
* This Abstract is published by the Society at 3 Hanover Square, London,
W., on the Tuesday following the date of Meeting to which it refers. It will
be issued, free of extra charge, to all Fellows who subscribe to the Publications,
along with the ‘ Proceedings’ ; butit may be obtained on the day of publication
at the price of Sixpence, or, if desired, sent post-free for the sum of Sir
Shillings per annum, payable in aryance.
22
with the last-named it was pointed out that the Cirrhitide, as
defined and limited by Dr. Giinther, with the addition of Haplo-
dactylus, form a very natural family.
Mr. Hamitron H. Drucs, F.LS., F.Z.8., read a short paper
«On some New and Little-known Hesperide from Tropical West
Africa,” which contained remarks on, and descriptions of, some
new forms of these Butterflies lately obtained by Mr. G. L.
Bates, F.Z.S., on the Ja River, Cameroons, and others from
Nigeria. New species of the genera Abantis, Acleros, Gorgyra,
Parnara, and Ceratrichia were described.
The next Meeting of the Society for Scientific Business will
be held on Tuesday, the 27th April, 1909, at half-past Hight
o'clock p.M., when the following communications will be
made ;—
1. Dr. T. A. Cuarman, F.Z.S.—A Review of the Species of the
Lepidopteran Genus Lycenopsis Feld. (Cyaniris auct. nec Dalm.)
on examination of the Male Ancillary Appendages.
2. KF. KE. Bepparp, M.A., F.R.S., F.Z.S.—On some Points in
the Structure of Galidia elegans, and on the Postcaval Vein in
Carnivora.
3. Dr. R. W. Snuretpr, C.M.Z.8.—On the Comparative
Osteology of the Passerine Bird Arachnothera magna.
Communications intended for the Scientific Meetings of the
ZOOLOGICAL Socrety or Lonpon should be addressed to
P. CHALMERS MITCHELL,
Secretary.
3 HANovER Square, Lonpon, W.
April 13th, 1909,
No. 70.
ABSTRACT OF THE PROCEEDINGS
ZOOLOGICAL SOCIETY OF LONDON,*
April 27th, 1909.
Prof. K. A. Mincutn, M.A., Vice-President,
in the Chair.
The SecrErary read a Report on the additions that had been
made to the Society’s Menagerie during the month of March
1909.
Mr. H. F. McSuHane exhibited a series of lantern-slides of
animals living in the Society’s Gardens.
Prof, K. A. Mincutn, M.A., V.P.Z.S., exhibited specimens of
a Tick which causes an African relapsing fever in man.
Mr. R. H. Burnu, F.Z.8., exhibited a series of specimens
from the Museum of the Royal College of Surgeons, showing
(1) the different mechanisms for the movement of the nictitating
membrane, (2) the coarse anatomy of the tapetum lucidum,
(3) the organ for elevating the eyes in the Plaice.
Mr. R. I. Pococn, F.LS., F.Z.S., exhibited photographs of the
two races of Burchell’s and Wahlberg’s Quagga recently imported
from Zululand by Herr Reiche.
A paper was presented by Dr. T. A. CHAapman, F.Z.S., entitled
‘“A Review of the Species of the Lepidopteran Genus Lyccenopsis
Feld. (Cyaniris auct. nec Dalm.) on examination of the Male
Ancillary Appendages.”
* This Abstract is published by the Society at 3 Hanover Square, London
W., on the Tuesday following the date of Meeting to which it refers. It will
be issued, free of extra charge, to all Fellows who subscribe to the Publications,
along with the ‘ Proceedings’; but it may be obtained on the day of publication
at the price of Sixpence, or, if desired, sent post-free for the sum of Sir
Shillings per annum, payable in adyance,
24
Mr. F. E. Bepparp, M.A., F.R.S., F.Z.8., read two communi-
cations: “On some Points in the Structure of Galidia elegans, and
on the Postcaval Vein in Carnivora,” and “On the Postcaval
Vein and its Branches in certain Mammals.”
A paper was received from Dr. R. W. Suurenpr, C.M.Z.S.,
“On the Comparative Osteology of the Passerine Bird Arachno-
thera magna,” the communication being based on material
supplied from the Prosectorium of the Society.
The next Meeting of the Society for Scientific Business will
be held on Tuesday, the 11th May, 1909, at half-past Hight
o'clock p.m., when the followmg communications will be
made :-—
1. Prof. Wm. RipcEeway, M.A.—
(a) On hitherto unrecorded Specimens of Hquus quagga.
(6) Differentiation of the Three Species of Zebras.
(c) On a Portion of a Fossil Jaw of one of the Kquide.
9. R. LypexKer.—On a new Race of Deer from Sze-chuen.
3. E. C. Cuuss, F.Z.8.—The Batrachians and Reptiles of
Matabeleland.
Communications intended for the Scientific Meetings of the
Zootocicay Socrery or Lonvon should be addressed to
P. CHALMERS MITCHELL,
Secretary.
3 Hanover Square, Lonpon, W.
May Ath, 1909.
THE ZOOLOGICAL SOCIETY OF LONDON.
NN OT
Tats Society was founded in 1826 by Sir Sramrorp Rarrtes,
Mr. J. Sasine, Mr. N. A. Vieors, and other eminent Naturalists,
for the advancement of Zoology and Animal Physiology, and for the
introduction of new and curious subjects of the Animal Kingdom,
and was incorporated by Royal Charter in 1829.
COUNCIL.
HIS GRACH THE DUKE OF BEDFORD, K.G., President.
Grorcze A. BoutEeneur, Ese.,
F.R.S., Vice-President.
Rost Braprorp,
Vice-
Pror. Joun
JWEIDES* IDSOhss dial tise
President.
Lr.-Cot. Sire R. Havetock
Cuartgs, K.C.V.O., M.D.
AtrreD H. Cocks, Esa., M.A.
Tue Rr. Hon. Kart or Cromer,
P.C., G.C.B.
Cuartes Drummonp, KEse.,
Treasurer.
Freperick GInterr, Hse.
F. DuCanze Gopman, KEsea.,
D.C.L., F.R.S., Vice-President.
Tue Margvis or Hamitron, M.P.
Sypyvey F, Harwer, Ese., M.A.,
F.R.S., Vice-President.
Stir Eomwunp G. Loprr, Br.
_E. G. B. Meave-Watpo, Ese.
| Pror. Epwarp ALFrep Mrncarn,
M.A., Vice-President.
| P. Coatuers MircHect, Ese.,
MEAS DSc neDi ans
Secretary.
W. R. Ocrtviz-Grant, Esa.
ALBERT Pam, Ksa.
OxprieLp Tomas, Esea., F.R.S.
Avsyn Truevor-Barryr, Ese,
M.A.
Epwarp G. Wappitove, Ilsa.
A. SuirH Woopwarp, Hse., LL.D.
E.R.S., Vice-President.
2,
The Society consists of Fellows, and Honorary, Foreign, and
Corresponding Members, elected according to the By-Laws. It
carries out the objects of its foundation by means of the collection
of living animals at Regent’s Park, by its Library at 3, Hanover
Square, W., and by its scientific publications.
The Office of the Society (3, Hanover Square), where all
communications should be sent, addressed to “The Secretary,” is
open from Ten till Five, except on Saturdays, when it closes at
Two P.M.
The Library, under the superintendence of Mr. F. H. Waterhouse,
is open daily at the above hours, except in September.
The Meetings of the Society for General Business are held at the
Office on the third Wednesday in every month of the year,
except in September and Octeber, at Five p.m. Commencing on °
January 20th, 1909, these Meetings will be held on the third
Wednesday of the month at 5 p.m.
The Meetings for Scientific Business are held at the Office twice
a month on Tuesdays, except in July, August, September, and
October, at half-past Hight o’cleck p.m.
The Anniversary Meeting is held on the 29th. of April, or the
nearest convenient day, at Four p.m.
The Gardens in the Regent’s Park are open daily from Nine o’clock
until Sunset. Mr. R. I. Pocock, F.LS., is the resident Superin-
tendent and Curator of Mammals and Reptiles. Mr. D. Seth-Smith
is Curator of Birds and Inspector of Works. The Prosectorium for
Anatomical and Pathological work at the Gardens is under the
charge of Mr. Frank E. Beddard, M.A., F.R.S., Prosector, assisted
by Mr. H. G. Plimmer, M.R.C.S., Pathologist to the Society.
TERMS FOR THE ADMISSION OF FELLOWS.
Fettows pay an Admission Fee of £5, and an annual Contri-
bution of £3, due on the Ist. of January, and payable in advance,
or a Composition of £45 in lieu thereof; the whole payment,
including the Admission Fee, being £50.
No person can become a Frttow until the Admission Fee and
First Annual Subscription have been paid, or the annual payments
have been compounded for.
Frriows elected after the 31st. of August are not liable for the
Subscription for the year in which they are elected.
3
PRIVILEGES OF FELLOWS.
Fettows have Personal Admission to the Gardens with Two
Companions daily, upon signing their names in the book at the
entrance gate.
‘The Wirz or Hustanp of a Fecxow can exercise these privileges
in the absence of the Fellow.
Every Fettow is entitled to receive annually 60 undated Green
Cards, and, when no specific instructions are received, the supply
will be sent in this form. If preferred, however, 20 Green Cards
may be exchanged for a book containing 2 Orders for each
Saturday * throughout the year. A similar book of Sunday Orders
may also be obtained in lieu of 20 Green Cards. A Green Card
may also be exchanged for 2 Buff Cards for the use of Children
under 12 years of age.
It is particularly requested that Fellows will sign every Ticket
before it goes out of their possession. Unsigned Tickets are not
available.
Green and Buff Tickets may be used on any day and in any year,
but in no case can two Children be admitted with one Adult
Ticket, or an Adult be admitted with two Children’s Tickets.
The annual supply of Tickets will be sent to each Fetrow on the
1st. of January in every year, upon filling up and returning the form
of Standing Order supplied to Fellows.
Frttows are not allowed to pass in friends on their written
Order or on presentation of their Visiting Cards.
Frniows are exempt from payment of the fee for Painting,
Sketching, and Photographing in the Society’s Gardens.
Frttows have the privilege of receiving the Society’s ordinary
Publications issued during the year upon payment of the additional
Subscription of One Guinea. This Subscription is due upon the
Ist. of January, and must be paid before the day of the Anniversary
Meeting, after which the privilege lapses. Fxtztows are likewise
entitled to purchase these Publications at 25 per cent. less than
the price charged to the public. A further reduction of 25 per
cent. is also made upon all purchases of Publications issued prior
to 1881, if above the value of Five Pounds.
Fettows also have the privilege of subscribing to the Annual
Volume of ‘ The Zoological Record,’ which gives a list of the Works
and Publications relating to Zoology in each year, for the sum of
* The Saturday Orders are not available if the Fellow introduces friends
personally on that day.
4
One Pound Ten Shillings. Separate divisions of volumes 39 to
42 can also be supplied. Full particulars of these publications can
be had on application to the Secretary.
Frettows may obtain a Transrerasre Iyory Ticker admitting
two persons, available throughout the whole period of Fellowship,
on payment of Ten Pounds in one sum, A second similar ticket
may be obtained on payment of a further sum of Twenty Pounds.
Any Fettow who intends to be absent from the United Kingdom
during the space of one year or more, may, upon giving to the
Secretary notice in writing, have his or her name placed upon the
“ dormant list,” and will be thereupon exempt from the payment of
the annual contribution during such absence.
Any Frttow, having paid all fees due to the Society, is at liberty
to withdraw his or her name upon giving notice in writing to the
Secretary.
Ladies or Gentlemen wishing to become Fellows of the Society
are requested to communicate with the undersigned.
P. CHALMERS MITCHELL,
Secretary.
8 Hanover Square, London, W.,
August Ist, 1909.
MEETINGS
OF THE
ZOOLOGICAL SOCIETY OF LONDON
FOR
SCIENTIFIC BUSINESS.
(AT 3 HANOVER SQUARE, W.)
1909.
Turspay, NovEMBER 9
09 ” 23
e DecemBeR 14
The Chair will be taken at half-past Light o'clock in the Evening
precisely.
LIST OF THE PUBLICATIONS
OF THE
ZOOLOGICAL SOCIETY OF LONDON,
Tue scientific publications of the Zoological Society of London
are of two kinds—“ Proceedings,” published in an octavo
form, and “‘ Transactions,” in quarto.
According to the present arrangements, the “ Proceedings”
contain not only notices of all business transacted at the scien-
tific meetings, but also all the papers read at such meetings
and recommended to be published in the ‘‘ Proceedings” by
the Committee of Publication. A large number of coloured
plates and engravings are issued in the volumes of the
“ Proceedings,” to illustrate the new or otherwise remark-
able species of animals described therein. Amongst such
illustrations, figures of the new or rare species acquired in a
living state for the Society’s Gardens are often given.
The “ Proceedings” for each year are issued in four parts,
on the first of the months of June, August, October, and
April, the part published in April completing the volume
for the last half of the preceding year. From January 1901
they have been issued as two half-yearly volumes.
The ‘‘ Transactions” contain such of the more important
communications made to the scientific meetings of the Society
as, on account of the nature of the plates required to illustrate
them, are better adapted for publication in the quarto form.
They are issued at irregular intervals.
Fellows and Corresponding Members, upon payment of
a Subscription of One Guinea defore the day of the Anni-
versary Meeting in each year, are entitled to receive the
Society’s Publications for the year. They are likewise
entitled to purchase the Publications of the Society at 25 per
cent. less than the price charged for them to the Public. A
further reduction of 25 per cent. is made upon purchases of
Publications issued prior to 1881, if they exceed the value of
five pounds.
Fellows also have the privilege of subscribing to the
Annual Volume of the Zoological Record for a sum of 30s.
(which includes cost of delivery), payable on the 1st. of July
in each year; but this privilege is forfeited unless the
subscription be paid before the Ist. of December following.
The following is a complete list of the publications of the
Society already issued.
TRANSACTIONS* OF THE ZOOLOGICAL SOCIETY OF LONDON,
Ato. 16 vols. and Index. ecole Price to the
ellows. Public.
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PROCEEDINGS OF THE COMMITTEE OF SCIENCE AND
CORRESPONDENCE OF THE ZOOLOGICAL SOCIETY OF
LONDON. 8vo. 2 vols. (Letterpress only). rice to Brice toithe
Teeth le! ISO Sales IO ChiOn GAbocomeones 5 AS 160.2 ee Osa
TI. 1832. Lith kee: erties As cea he ates
99
PROCEEDINGS OF THE ZOOLOGICAL SOCIETY OF LONDON.
8vo. 15 vols. (Letterpress only) and Index. (First Series.)
Price to Price to the Price to Price to the
Fellows. Public. Fellows. Public.
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LISTS OF THE ANIMALS IN THE SOCIETY’S GARDENS.
List of the Vertebrated Animals now or lately Living in the Gardens
of the Zoological Society of London. (Kighth Edition.) 8vo.
1883. Cloth, 4s. 6d.
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THE OFFICIAL ILLUSTRATED GARDEN GUIDE—7th Edition
—through this Society’s Gardens, with (1) a Street Map,
showing a direct route to the ‘‘ Zoo” from all parts of London
and Suburbs ; (2) a Plan of the Grounds, showing at a glance
the location of the animals ; (3) a short description of some of
the principal animals in the Collection (now containing about
3300 specimens), together with 50 Photographic Illustrations
and Index, may now be obtained at the Society’s Office,
3, Hanover Square, W., or at the Society’s Gardens in
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P. CHALMERS MITCHELL,
Secretary.
3, Hanover Square, London, W..,
August Ist, 1909.
These publications may be obtained at the Socrery’s Orrice
(3, Hanover Square, W.), at Messrs. Lonamans’ (Paternoster Row,
E.C.), or through any bookseller.
ZOOLOGICAL SOCIETY OF LONDON.
THE ZOOLOGICAL RECORD.
ie object of the ZootogicaL Recorp is to give, by means of an
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the year preceding the issue of the Volume, together with full
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manner as to serve aS an Index to the literature of Zoology in all
parts of the globe, and thus to form a repertory that will retain its
value for the Student in future years.
The ‘ Zoological Record’ having been amalgamated with the
International Catalogue of Scientific Literature, Zoology, Volumes
from 43 onwards can only be obtained now from Messrs. Harrison
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Divisions of the ‘ Zoological Record’ of Vols. 39 to 42 can be
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[P. i, ©,
SEPARATE DIVISIONS OF THE ZOOLOGICAL RECORD.
Divisions of the ‘Zoological Record,’ Vols. 39-42, containing
the literature of the years 1902-1905, may be obtained separately
as follows :—
Con
bo %
S
)
Pr
List of abbreviations of journals, etc.
Special Records, viz. :—
I. General Subjects ..
II. Mammalia
III. Aves ie ees lee
IV. Reptilia and Batrachia. .
V. Pisces
VI. Tunicata
VII. Mollusca 5
VIII. Brachiopoda..- ..
IX. Bryozoa
X. Crustacea
XI. Arachnida
XII. Myriopoda
XIII. Insecta
XIV. Echinoderma
XV. Vermes ..
XVI. Celenterata ..
XVII. Spongie
XVIII. Protozoa
=
NONrFWwWNrHE WY NDH KH Be wd Db bw
é
S 29 QF oe One oo SoS EE @ CO &® eo
bo
Index of new names of genera and subgenera. 7
Divisions from Vol. 43 onwards are now supplied by Messrs.
Harrison & Sons, 46 St. Martin’s Lane, London, W.C.
P. CHALMERS MITCHELL,
Secretary.
8 Hanover Square, Lonpon, W.
August 1st, 1909.
ConTENTS (continued).
April 6, 1909.
Mr. George Jennison, Exhibition of some fertilized eggs from a pair of Seba Pythons.... 692
Dr. R. T. Leiper, F.Z.S. Exhibition of a distorted Elephant’s tusk and a malformed canine
HAOUMY Oli ae ELI OPGHANMEIS ss. cin -us 5 =o se ss ae se sessing semis Foe oOOwowS Co AestT ++ 390
Mr. E. T. Newton, F.R.S., F.Z.S. Exhibition of a metatarsal bone of an Ox, showing the
marks of gnawing by rodents... sth ot lvoe, ee ee a eee he 395
1. Description of a new Form of Ratel (Mellivora) from Sierra Leone, with Notes upon the
described African Forms of this Genus. By R. I. Pococg, F.LS., FZ. 8., Superintendent
mugune: Grandens.sc (blate Nl ys se cw oreteaieieasa eats seta orors ais Gis Hiner oN OCS Scene og4
2. Notes on an Ichthyosporidian causing a Fatal Disease in Sea-Trout. By Murier
Rogpertson, Carnegie Research Fellow. (Plates LXII.-LXIV.)..................-. 399
3. A Collection of Fishes made by Dr. C. W. Andrews, F.R.S., at Christmas Island. By
@2 Tarn Reon, M.A., F.Z.S8. (Plates DXV. & LXVI.) 2. cee wee cen cscs ceca se 405
4, On some New and Little-known Hesperiide from a ae West Africa. fe HAMILTON
H. Druce, F.LS., F.Z.8. (Plate LXVII.) .. Srasiersvar cledeliys \e'al ciate dhol aoa cetagtne st ayiobeiays 406
April 27, 1909.
The Secretary. Report on the Additions to the Society's Menagerie during the month of
Marcell O Dire eiatehien sy are lrarsteneiate a\lcian severemeye Wve ersias ensv rs) = SO G6 botlaeose.ocub SoInaOCaes 414
Mr. H. F. McShane. Exhibition of lantern-slides of animals living in the Society’s
Grandlentsiniem sie se Ob D SGI8 BOG GA eie be aren CL En Oar RCI Ere Mie Eben ota cirericte pein Ge COiar 414
Prof. KE. A. Minchin, M.A., V.P.Z.S. Exhibition of the eggs and living specimens of the
Tick, Ornithodoros moubata Whey eS Soocece sey staat eeNeee crestor Sa cobmsagoo nod ec -. 414
Mr. R. H. Burne, F.Z.8. Exhibition of Anatomical mechanisms of the eyes of some
AVGRUS PES inipicte Glted cuclsuoion ee ociunlnlolnttadine 010. MgO OapInc bone ne bo weindion Docc 414
Mr. R. I. Pocock, F.L.S., F.Z.S. Exhibition of, and remarks upon, the photographs of
two Quageas............ sietaises calute edatader ind oenemonos COO cobmeDaO re aislcichatchs cticek aeuase 415
‘|. A Review of the Species of the Lepidopterous Genus Lycenopsis Feld. (Cyaniris auct.
nec Dalm.) on examination of the male Ancillary Appendages, By T. Avcurnon
Carman. M.D., F.ZS. . SED Omen ee Oh Ob hin oo Bhi tahoe ae eo sine Gopecnercn wll!
. On some Points in the Structure of Galidia elegans, and on the Postcaval Vein in
Carnivores By Franx E. Beppanp, M.A., F.R.S., F.Z.S., Prosector to the Society.... 477
3. On the Postcaval Vein and its Branches in certain Mammals. By Frank EB, Bepparp
M.A., F.R.S., F.Z.S., Prosector to the Society
Sorrel wis; 4 sVare wrote ty senate Gs ale erallcre\ afapatee SOG)
4. On the Comparative Osteology of the Passerine Bird Arachnothera ee By R. W.
SHUFELDD eM OiMEZ: Ss (blaterisXevallls bene gas | cerca ae OSs tiga acer
Minlepage: een ws co Dosen cseiaieariaeater ex's eitey sbesenstc Fe Rey ct Mieke causles cei e if ta ee Rook te noe EN oe ee i
_ List of Council and Officers ii
List of Contents ........ Pere wide Ma rope ee ARNON Son vere un eper otal ht Ti
“AlphabeincalMbistrat) Contributors sisi s Gen baler aia) tere Winey 20 one th Ps rece ix
LETS ASS 2 0 at 2 a ae Re arte in) ri teat ee pene emporio le xvii
Paebiol Text-tia tines ie wmmetarcts i) aj.) uitl ow ncsiss sos lei ee ae ene ee ee moe RES. 50 babs
List of New Generic Terms . Xx
RTCLe KM 4, ss aes
1909, pp. 201-544,
Plate Page
sat Shells of Patella wulgata, Boe. +. tee epee
XXXIV. } Development of Agrionid Dragonflies ..............-065 253
XXXYV.)
XXXVI. |
XXXVIT. } Goniodes tetraonis .....6... Fo odin. Sol Mee Meee \
XXXVIIL | inet
XXXIX. )
XL. .Goniodes tetracnis and Menopon pallescens i emiisie. ts |
MI WGoniodestandeNirmus) Sener: seniemedsee ce eee eee |
AD ING is COMenGUUS: * Lois ok se hates La tie weteie cs ne en ie ee
= ALIN a rer is r
XLIV. | WUONO MY LE CAG OPOGLA s.r An, We eee tates +)-) Som nad eee
ae } Seatophag a SiercOndnig ye celal = late. 3 he eiate. eee ea er
XLVII. Ceratophyllus gallinul@ 6... -6cc cece eee ete eens ee)
XEVIII. \
a " Trichostrongyus Per gracias scias osae cvae conta ec sete nue \
ut Trichostrongylus pergractiis and Trichosoma longicolle .... ae
Se \ UEP iChOSONLANONGIGONE exec ert rale in\e staloverelsiay icicle een etka |
LV. Contents of alimentary canal of Grouse ............... )
TVA, “Dacaitied rogauitn a 0. 2s efor aie ste sioreints Hela « «'eiciuneias aeiies |
LVIL. Davainea urogalli and Hymenolepis microps . ie See i, |
LVIIL. Hymenolepis mtcrops...- sees cscceccenseerrvseecssornce $ 351
TENG ODOT CROFT ALL vinheiw o secte slave ol sraiesloe etl) ct ene eee |
LX. Davainea urogalli and Hymenolepis microps ............--
LXI. Mellivora signata .......... aiataepiarepSichedt-retaoese cova eeent Ae cae nan 394
LXIT.
ipxcalane | Iehthyosporidians Of-Néd- Trout sem ss ekbene Ce Cee 399
LXIV :
LXV. Parupeneus andrews 1s. seen ee ne ce ee ne ence nese
LXVI. 1. Blennius atrocinctus, 2, BL. nativitatis. 3. Salarias 4
caudofasciatus. 4. S. natalis. 5. S. melanosoma. 408
G. Cirrhites MUTTAYE co cnc vanesevensscceven cn cicoe ae
LXVII. New and Little known W. African ee sin often Nemes 406 -
LXVIII. Skeleton of Arachnothera magna ......- vid’. Wier CeO
NOTICE.
The ‘ Proceedings’ for the year are issued in four parts, paged consecutively,
so that the complete reference is now P. Z. 8. 1909, p. .. . The Distribution
is as follows:—
Papers read in January and February, issued in June.
x“ » March and April, » 9», August.
EF » May and June, Pee October:
i », November and December, ,, ,, April.
Proceedings,’ 1909, pp. 1-200, were published on May 24th, 1209.
The Abstracts of the papers read at the Scientific Meetings in
March and April are contained in this Part.
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